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Plant Species Biology (2013) 28, 130–137 doi: 10.1111/j.1442-1984.2012.00366.x

Strategies of plant establishment of two Cerrado species:

Byrsonima basiloba Juss. (Malpighiaceae) and Eugenia
dysenterica Mart. ex DC (Myrtaceae) psbi_366 130..137

CONCEIÇÃO E. S. SILVEIRA, DARIO PALHARES, LUIZ A. R. PEREIRA, KARINA B. D. PEREIRA and

FERNANDA A. B. SILVA
Laboratory of Plant Anatomy, Department of Botany, University of Brasília, Campus Darcy Ribeiro, Postcode 70910-900,
Brasília, DF, Brazil

Abstract
This study investigates the initial development of two Cerrado species, Eugenia dysen-
terica and Byrsonima basiloba. Both species have high ethnobotanical importance and are
widely used as a food source as well as in folk medicine. Eugenia dysenterica presented
hypogeous cryptocotylar germination and the cotyledons were rich in reserve material.
This plant species showed quick root development and after the shoot had produced the
first two pairs of eophyls, the seedling went into a latent state for at least 1 year. Byr-
sonima basiloba had a hard endocarp, which had to be broken to facilitate germination.
This species showed epigeous phanerocotylar germination. Similarly to E. dysenterica,
B. basiloba roots also developed rapidly; however, the aerial part grew continuously and
did not show a latent period. Moreover, within the first 6 months of cultivation B. basi-
loba developed a xylopodium. The strategies for seed dispersal and breakage of seed
dormancy were different between the species, but their initial growth showed several
similarities.
Keywords: clay, cultivation, fire, seedling, soil.
Received 15 June 2011; revision received 29 September 2011; accepted 29 October 2011

Introduction sclerified endocarp, which surrounds the seeds. The

endocarp must be broken to allow the seeds to absorb
In general, the Cerrado is a harsh environment and young
water and germinate. Under natural environmental con-
developing plants have to face the dry season and wild
ditions, it is inferred that the seeds only germinate after
fires, as well as the poor acid soils of this savannic vegeta-
fire, which is the main agent that breaks the hard
tion (Franco 2004). Therefore, Cerrado plants present
endocarp (Araújo 1994).
many surviving strategies that encompass the timing of
With respect to Eugenia, this genus is found throughout
germination and sprouting, fast root growth, fire-resistant
tropical and subtropical America. The Cerrado has
structures and the development of resprouting structures,
approximately 15 species, and E. dysenterica, also known
such as xylopodia and budding roots (Palhares & Silveira
as cagaita, is the one with the largest phytosociological
2007).
predominance (Almeida et al. 1998). It is a tree with thick
The genus Byrsonima is distinct from other Malpighi-
and fire-insulating bark and is characterized by a large
aceae because it produces pulpous and edible fruits,
production of edible fruits that have been widely used in
which are popularly known as murici (Barroso 1991). In
juices, ice creams and jellies (Proença & Gibbs 1994; Silva
the Cerrado, there are approximately 35 species of Byr-
et al. 2001; Palhares 2003; Souza et al. 2008; Pina et al. 2009).
sonima and B. basiloba has the largest fruits (Ferreira 1973).
As both species have been targeted for domestication
This plant is a shrub that generally bears fruit all year
owing to their multiple uses for pharmaceutical and food
long. The fruit is a drupe, possessing a juicy pulp and a
source purposes, the present study investigates the initial
morphological development of B. basiloba and E. dysen-
Correspondence: Dario Palhares terica. As both species are relatively common as phytoso-
Email: dariompm@unb.br ciologically dominant species and because they come

© 2012 The Society for the Study of Species Biology

 E S TA B L I S H M E N T O F B Y R S O N I M A A N D E U G E N I A 131

from different families, the strategies they present in their For both species, the stages of germination and devel-
first years of life can highlight the necessities that a plant opment were observed daily in the laboratory and
may need to possess to establish in such an environment. samples were taken from the glasshouse and described.
Each described stage corresponds to 10 individuals.

Materials and methods

Seeds of E. dysenterica (n = 70) and B. basiloba (n = 436)
were collected in Cerrado areas in the vicinity of Brasília
from mature and healthy plants (4–6 m in height and
25–30 cm trunk circumference for E. dysenterica and 2–4 m
in height and 10–14 cm trunk circumference for B. basi-
loba). Voucher specimens were deposited at the herbarium
of the University of Brasilia (UB) under the registration
numbers UB 11689 (E. dysenterica) and UB 11690 (B.
basiloba).
Two sets of E. dysenterica seeds were put to germinate in
either agar–agar solid medium or in soil mix. A small
puncture of 3–5 mm was made in the testa as suggested
by Rizzini (1970) because this procedure enhanced germi-
nation rates. The group of seeds germinated in agar–agar
was disinfected in 70% ethanol for 5 min and in commer-
cial bleach (2% active chlorine) for 40 min. Subsequently,
the seeds were rinsed three times in water and inoculated
in test tubes containing sterilized agar water (7 g/L). The
culture conditions were: 25 ⫾ 2°C, 80% relative humidity,
12 h photoperiod and a light intensity of 41 mmol/m2/s.
This set of seeds was used to study the initial seedling
development of E. dysenterica seedlings for 30 days. The
purpose of germinating the seeds in agar–agar was to
facilitate the initial development of the root system.
The other group of seeds was cultivated in plastic cups
containing a mixture of vermiculite, peat-based substrate
and red latosol (1:1:1) for approximately 1 year in a green-
house. This set of plants was used to study the develop-
mental processes of older plants of E. dysenterica.
Byrsonima basiloba fruits had the pulp extracted and the
endocarps were washed and brushed under tap water.
After this, the endocarp was dried and broken to remove
the seeds. Two sets of seeds were prepared and cultivated
as described for E. dysenterica.
Results
Eugenia dysenterica
Eugenia dysenterica seeds are either round or elliptical in
shape with a diameter of approximately 2 cm, a coriaceous
tegument and cotyledons with high reserve content. The
seeds show hypogeous cryptocotylar germination.
Most E. dysenterica seeds germinated between the 7th
and 19th days. In the glasshouse, the first signs of the
germination process occurred at 35 days of cultivation,
when the epicotyl emerged from the substrate. On the
38th day, 96% of the seeds had already germinated. After
82 days, there were 59 well-established seeds (88% of the
germinated seeds) and five plants (8% of the seedlings)
presented abnormal characteristics, such as root death,
the absence of the hypocotyl and non-specific malforma-
tion of the roots and aerial parts.
Table 1 summarizes the post-seminal phases and
Figure 1 illustrates the developmental stages of the seed-
lings. The first structure to appear is the protruded
hypocotyl-root axis right after the rupture of the seed coat.
At this moment, at the micropyle region, the seed coat
broke, but remained attached to the cotyledons for the
entire period of observation (Fig. 1a–f). At the beginning,
the hypocotyl and primary root had no clear separation
(Fig. 1a,b). As the developmental process continued, the
primary root became clearly distinguished from the hypo-
cotyls; this was observed from days 25 to 30 of cultivation
(Fig. 1c,d). Initially the hypocotyl was horizontal (Fig. 1c),
which placed both the superior part and the root system
in a perpendicular position to the hypocotyl (Fig. 1d,e).
This caused a longitudinal discontinuity between the epi-
cotyl and the root (Fig. 1c). At approximately day 60, the
hypocotyl had already tilted and assumed an upright

Table 1 Chronological phases of the

initial development of seedlings of Eugenia Days after
dysenterica Morphological milestone imbibition Figure 1

Protrusion of the primary root 7–19 a

Emission of the primary root 21–28 b
Emission of the epicotyl 26–37 c
Elongation of the epicotyl and formation of 30–40 d
the cataphylls
Emission of lateral roots ~45 d
Emission of the first pair of eophyls 66–123 e
Emission of the second pair of eophyls 78–146 f
Beginning of the latent phase ~180–700 (at least) Not shown

Plant Species Biology 28, 130–137 © 2012 The Society for the Study of Species Biology
132 C . E . S . S I LV E I R A E T A L .

Fig. 1 Aspects of the germination, seedling and young plant of Eugenia dysenterica Mart. ex DC. (a–e) Seedling at (a) 10 days, (b) 25 days,
(c) 30 days, (d) 45 days and (e) 70 days. (f) Young plant at 90 days. AB, apical bud; Ca, cataphyll; Co, cotyledon; Ep, epicotyl; Eo, eophyl;
Hp, hypocotyls; LR, lateral root; PR, primary root; T, testa.

© 2012 The Society for the Study of Species Biology Plant Species Biology 28, 130–137
 E S TA B L I S H M E N T O F B Y R S O N I M A A N D E U G E N I A
position and both the epicotyl and root system continued
longitudinally (Fig. 1d–f). In addition, the axial root was
cylindrical, hairless, tortuous, reddish and thicker at the
hypocotyl region (Fig. 1e,f). The primary root grew con-
tinually, and the first lateral roots appeared on the 45th
day of development (Fig. 1d–f).
The aerial parts of E. dysenterica seedlings took longer
than the roots to develop and mostly appeared after
approximately 30 days of cultivation (Table 1). The epicotyl
was initially reddish. After 2–3 months it turned green and
then slowly it turned brown. The stem continued to grow
and the first pair of cataphylls was observed around day 30
of observation (Table 1). Five to eight pairs of cataphylls
were emitted before the first eophyls (Fig. 1d,e). The phyl-
lotaxy of the cataphyls was opposite-decussate.
It is also noteworthy that the epicotyl began to elongate
only after the primary root had reached 3.5–4.0 cm in
length, after 3 months of cultivation. The internode of the
first pair of eophyls began to elongate, and the second pair
of eophyls appeared, making the phyllotaxy crossly
opposed as in the mature plant. The total length of the
plantlet was approximately 25 cm, with 18 cm of root and
7 cm of aerial growth. In general, 6-month-old seedlings
presented two pairs of eophyls (Fig. 1f). The eophyls had
a short petiole and were chartaceous, hairless and
lanceolate-elliptic, with smooth margins and a distinct
midrib. The eophyls and the adult plant leaves were mor-
phologically similar. Furthermore, in some plants the first
two pairs of eophyls senesced and fell and new ones were
formed. After the formation of the second pair of eophyls
the growth of the seedlings slowed down, and at approxi-
mately 6 months of age they entered a latent phase with
no noticeable shoot growth, which lasted for at least
2 years. During this period, the eophyls occasionally died
and two new pairs appeared, but the final length of the
aerial portion increased by only a few more centimeters.
Byrsonima basiloba
Byrsonima basiloba seeds were roundish and approxi-
mately 5 mm in diameter. Each endocarp had three loculi
with one seed in each and, in general, each fruit had only
one viable seed. The seed had a pointy end at the radicle
region with a membranaceous, soft and brown testa. In
contrast to E. dysenterica, B. basiloba had epigeous phan-
erocotylar germination. A total of 86% of the seeds germi-
nated. Approximately 15% of the plantlets showed
abnormal characteristics, such as incomplete expansion of
the cotyledons, the formation of a callus in the epicotyl
and the formation of adventitious roots on the cotyledons.
Table 2 shows the chronology of some post-seminal
developmental milestones and Figure 2 illustrates several
phases of seedling development. Germination started 1–3
days after imbibition (Fig. 2a,b). In most cases, the break-
Plant Species Biology 28, 130–137
133
Table 2 Chronological phases of the initial development of seed-
lings of Byrsonima basiloba
Days after
Morphological milestone imbibition Figure 2
Protrusion of the primary root 1–3 b
Protrusion of the cotyledons 4–7 c
Expansion of the cotyledons/green color 8–11 d
Emission of the primary root 11–15 c,d
Emission of lateral roots 14–30 e
Expansion of the hypocotyl 25–30 e,f
Emission of the first eophyls 30–40 f
Emission of the 2nd/4th pair of eophyls 50–60 g
ing of the seed coat took place in the region of the radicle,
resulting in the protrusion of the primary root through the
newly formed gap (Fig. 2a). However, in a few seeds, the
seed coat broke in the cotyledonary region and the coty-
ledons were the first organs to appear. On the 4th day, the
cotyledons, which were colorless and partially circinate
(Fig. 2c,d), started to turn green. Fully expanded cotyle-
dons were elliptical (Fig. 2e), hairless, dark green and
opposite, with a decurrent base, obtuse apex, entire
margins and a pronounced and symmetrical midrib.
At approximately the 15th day of inoculation, subse-
quent to the initial elongation of the primary root, the
cotyledons initiated the expansion and unfolding pro-
cesses, which continued until the 30th day (Fig. 2d,e). The
lateral roots developed only after the cotyledons had fully
expanded and continued to show up throughout root
development. In 30-day-old seedlings, the primary root
was axial, approximately 5 cm long, and both the primary
and lateral roots were slightly waved (Fig. 2e). Root hairs
were observed in all roots. In those plants in which the
epicotyl protruded before the primary root, lateral roots
also appeared after the cotyledons had completed their
expansion.
After the cotyledons had completed their expansion,
the epicotyl started to grow, and the first eophyls
appeared at the lower region of the epicotyl. After
2 months, the seedlings were approximately 8 cm long,
and two-thirds of their total seedling length comprised
the root system (Fig. 2f). In addition, the seedlings had
2–4 pairs of eophyls (Fig. 2f,g). The epicotyl internodes
were roundish, densely hairy and pliable.
The eophyls were sessile or subsessile, with entire
margins, intrapetiolar stipules and opposite-decussate
phyllotaxy. The first pair was ovate, with acute apex,
obtuse base and trichomes only in the midrib and mar-
ginal veins. The second pair was much larger than the first
pair, with an elliptical shape, an acute apex and an obtuse
to roundish base, with malpighiaceous trichomes. More-
over, the second pair was glabrescent on the adaxial side
and densely pilous on the abaxial side.
© 2012 The Society for the Study of Species Biology
134 C . E . S . S I LV E I R A E T A L .

Fig. 2 Germination of the seed and development of a seedling and young plant of Byrsonima basiloba Juss in relation to the phases of
Table 2. (a,b) Third day of cultivation. (a) Rupture of the tegument by the cotyledons. (b) Appearance of the primary root. (c) The
cotyledons start to elongate on the 7th day. (d) The cotyledons are more elongated and the primary root shows continuous growth on the
15th day. (e) Well-expanded cotyledons and the hypocotyl expands on the 30th day. (f) First pair of expanded eophyls and primary root
with many secondary roots on the 40th day. (g) On the 60th day, two to four pairs of eophyls with crossly opposed phyllotaxy, thickening
of the hypocotyl and some tertiary roots. At right: 2-year-old young plant showing a xylopodium, a well-developed root system and a
straight stem without branches. AB, apical bud; Co, cotyledon; Eo1, first pair of eophyls; Eo2, second pair of eophyls; Hp, hypocotyl;
LR, lateral root; PR, primary root; T, testa; Xy, xylopodium.

© 2012 The Society for the Study of Species Biology Plant Species Biology 28, 130–137
 E S TA B L I S H M E N T O F B Y R S O N I M A A N D E U G E N I A
The primary root had a lateral and oblique position
in relation to the hypocotyl base (Fig. 2g). This region,
termed the collet, possesses a small appendix below the
region where the primary root appears. The lower region
of the hypocotyl was thicker and it was in this region that
the xylopodium usually developed (Fig. 2g,h). Figure 2h
shows a 2-year-old plant with a well-developed xylopo-
dium and root system. In addition, we can see that the
cotyledons and eophyls had fallen and compared with
the eophyls the mature leaves were morphologically
different. These leaves were either sessile or subsessile,
stipulated with an auriculate base and an obtuse shape
(Fig. 2h).
Discussion
Germination
Eugenia dysenterica Other studies on the germination of
E. dysenterica have recorded a germination rate varying
from 9.5 to 95% (Naves et al. 1992; Oga et al. 1992). This
plant presents an atypical dormancy. Rizzini (1970)
carried out several experiments on the germination of
E. dysenterica seeds and showed that fresh seeds quickly
and freely absorbed water and oxygen. However, the
tegument contains a large number of phenolic com-
pounds as evidenced by histochemical tests with iron
chlorine. When the tegument is removed the seed quickly
germinates; however, when just a small wound is made
on the tegument, germination rates rise to up 95% and
histochemical tests no longer show phenolic compounds.
However, scarification is only effective for recently col-
lected seeds owing to the fact that they are recalcitrant and
rapidly lose their viability when stored for more than
2 months (Cunha 1986; Farias-Neto et al. 1991; Andrade
et al. 2003).
Baskin and Baskin (2004) offered a general classification
of seed dormancy and Linkies et al. (2010) reviewed seed
dormancy from an evolutionary point of view. In this way,
E. dysenterica presents mild physiological dormancy, but
it does not fully match the Class A of Baskin and Baskin
(2004): Class A is related to gibberelin, temperature, iso-
lation of the embryo and a dry period, which is not exactly
the case for E. dysenterica. In addition, Linkies et al. (2010)
called attention to a specific type of dormancy that can be
broken by the removal of the testa and/or the endosperm.
Eugenia dysenterica is again a particular case because the
experiments of Rizzini (1970) highlighted that the mecha-
nisms of seed dormancy are related to phenolic com-
pounds that are somehow destroyed after a small wound
in the testa.
Eugenia dysenterica seed dispersion takes place just
before the beginning of the wet season (Sano et al. 1995;
Souza et al. 2008), and Brito et al. (2003) found that in the
Plant Species Biology 28, 130–137
135
fields seedlings are already established within 6 months
from the beginning of the rains.
With regard to the morphological aspects of the young
plants, our results are also consistent with those of
Andrade et al. (2003) and Duarte et al. (2006).
Byrsonima basiloba With respect to the Malpighiaceae
family, there are very few studies that have examined seed
morphology and germination, with the exception of Souto
and Oliveira (2005, 2008). In this family, seeds vary widely
in terms of their shape; however, they generally have
membranaceous teguments as observed in B. basiloba and
other species of the genus such as Byrsonima intermedia
(Souto & Oliveira 2005). Byrsonima basiloba presented
epigeous phanerocotylar germination, which is quite
common in other Malpighiaceae members, such as Byr-
sonima coriaceae (Duke 1965, 1969), as well as in Banisteri-
opsis (Gates 1982). However, cryptocotylar germination
may also be found in this family as observed in Heterop-
terys laurifolia (Duke 1965).
In general, plant reproduction is under the influence of
several environmental cues, such as drought, rainfall and
temperature; in Cerrado vegetation, fire is also an impor-
tant component and some species with covered seeds
require fire to break the cover and release the seeds
(Hoffman 1998; Silva et al. 2001). However, B. basiloba seed
production and dispersion does not appear to have a
pattern associated with such environmental stimuli
because this species bears fruit all year long.
In B. basiloba, the tegument was adnate to the
endocarp, which was hard and impermeable. This type
of adnation has also been observed in other species
of the genus such as Byrsonima verbascifolia (Almeida
et al. 1998). Furthermore, because of its structural char-
acteristics the endocarp was a severe obstacle for seed
germination. Our data showed that the endocarp must
be broken in order for the seed to germinate. Although
there are no reports on the germination rate of this
species in natural conditions, the rigidity and imper-
meability of the seed–endocarp complex may likely
cause germination hindrance. In addition, B. basiloba
seeds are recalcitrant and thereby have lower resistance
to dehydration and consequently limited viability after
dispersal.
The endocarp–seed complex has been considered the
true diaspore of B. basiloba (Barros 1992). According to the
classification of dormancy of Baskin and Baskin (2004),
this plant could be classified as Class D: physical dor-
mancy. However, this classification takes the seeds into
consideration, which is not totally what happens in B. basi-
loba: the seeds germinate easily, but the fruit is the hard
and impermeable structure.
Thus, according to Finch-Savage and Leubner-Metzger
(2006), the seeds of B. basiloba are not strictly dormant
© 2012 The Society for the Study of Species Biology
136 C . E . S . S I LV E I R A E T A L .
because development of the embryo quickly starts after
imbibition at the physiological temperature.
It is clear that B. basiloba has a different reproductive
strategy than E. dysenterica. Although E. dysenterica waited
for the wet season to germinate, B. basiloba presented a
highly resistant endocarp that blocks water uptake as well
as gas exchange, and thereby the germination process.
Like other Byrsonima, such as B. intermedia (Souto &
Oliveira 2005), the tegument of B. basiloba was thin and
highly permeable to water. In addition, seeds with intact
embryos germinated almost immediately after they had
been exposed to water. Therefore, the presence of the
endocarp is an important hindrance to the germination of
this species in nature. Byrsonima basiloba is dependent on
dispersers, such as Rhea Americana (Faria et al. 2009), and
there is no direct association with specific environmental
stimuli. Therefore, the reproductive success of this plant
species might be dependent on other factors.
It is important to note that sprouting structures, such
as xylopodia, sprouting roots, tubers and others, are
common in Cerrado plants (Palhares et al. 2006). The
xylopodium in B. basiloba plants developed very early. To
the best of our knowledge there is no study that reports
this structure in this plant. This fact is of great ecological
relevance because in the natural environment this plant
has no protection against wildfires, and the aerial parts of
these plants are completely burned during this type of
event. However, field observations have shown that
B. basiloba regenerates after fire.
Seedling development and establishment
Species from different families tend to show different
manifestations of seed dormancy; there were several
similar strategies for the initial growth between the
species studied.
The formation of abnormal plantlets is common in
many species and both species presented approximately
the same percentage of abnormal plantlets. However,
the nature of the problem differed: E. dysenterica was
characterized by the death of organs, probably related to
excessive oxidation of phenolic compounds, whereas
B. basiloba showed problems in the growth and differen-
tiation of organs. In addition, polyembryony and subse-
quent irregular plantlet development was observed in
B. basiloba, but not in E. dysenterica (Salomao & Allem
2001).
The species examined in the present study have distinc-
tive strategies for seed dispersal: E. dysenterica fruits at the
beginning of the rainy season, in such a way that at 180
days the seedlings are struggling in their first dry season,
whereas B. basiloba, a shrub with a xylopodium, releases
seeds almost all year round and shows continuous
growth both in aerial and subterranean organs.
© 2012 The Society for the Study of Species Biology
Our study showed that E. dysenterica seedlings devel-
oped their root system prior to the shoot, whereas B. basi-
loba quickly develops the aerial portion. Although
E. dysenterica seedlings presented a 2-year-long latent
phase, after 2 years of growth B. basiloba was a 1-m tall
shrub. It is generally thought that E. dysenterica grows
downwards (roots) for five or more years and only after
that does it start to show significant growth of the stem (C.
Silveira, field observations, 2001). To date, long-term
studies on seedling and plant development of Cerrado
species remain an unexplored and challenging subject
(Duarte et al. 2006; Zaidan & Carrera 2008).
It is clear that E. dysenterica and B. basiloba show differ-
ent strategies for seed germination, which may directly
be associated with the seed structure. Eugenia dysenterica
had larger seeds, whose cotyledons had a relatively large
nutritional reservoir, whereas the cotyledons of B. basiloba
quickly emerged from the soil, allowing an immediate
initiation of photosynthesis. Nevertheless, both species
prioritized the growth of the subterranean system. The
growth rate of the roots and the physical structure of
Cerrado soils influence the ability to uptake water and
nutrients (Gross et al. 1992). Therefore, early ramification
of the root together with rapid vertical growth is an
adaptative strategy for survival; open and oligotrophic
fast root growth may be considered to be an important
strategy for survival in oligotrophic soils such as those of
the Cerrado (Moreira & Klink 2000).
The Cerrado vegetation as a whole is composed of a
considerable subterranean biomass, which exceeds that of
the aerial biomass. Thus, the ecological interaction of the
young plants with the soil, in the sense of protection
against fire and the ability to reach the deeper humid soil
layers, is an important feature that enables the plants to
overcome the selective pressure exerted by the Cerrado
environment, which determines the native species of this
biome.
Acknowledgments
We thank the Coordination for the Improvement of
Higher Level Personnel (CAPES), the Brazilian Council of
Research (CNPq) and the Foundation for Scientific and
Technological Enterprises (FINATEC) for their financial
support of this research. Thanks also to Neil Hume Wilson
for reviewing our English.
References
Almeida S. P., Proença C. E. B., Sano S. M. & Ribeiro J. F. (1998)
Cerrado: espécies vegetais úteis. Embrapa-CPAC, Planaltina-DF.
Andrade A. C. S., Cunha R., Souza A. F., Reis R. B. & Almeida J.
K. (2003) Physiological and morphological aspects of seed
viability of a neotropical savannah tree, Eugenia dysenterica
DC. Seed Science and Technology 31 (1): 125–137.
Plant Species Biology 28, 130–137
 E S TA B L I S H M E N T O F B Y R S O N I M A A N D E U G E N I A
Araújo A. R. B. (1994) Morfologia de frutos, sementes e plântulas, tipo
e aspecto da germinação de algumas espécies de malpighiaceae.
Unicamp, Campinas.
Barros M. A. (1992) Phenology of flowering, reproductive strate-
gies and pollination of sympatric species from the genus
Byrsonima (Malpighiaceae). Revista Brasileira de Biologia 52:
343–353.
Barroso G. M. (1991) Sistemática de angiospermas do brasil. UFV,
Viçosa.
Baskin J. M. & Baskin C. C. (2004) A classification system for seed
dormancy. Seed Science Research 14: 1–16.
Brito M. A., Pereira E. B. C., Pereira A. V. & Ribeiro J. F. (2003)
Cagaita: biologia e manejo. Embrapa-Cerrados, Brasília.
Cunha M. C. L. (1986) Estudo de preservação do poder germinativo de
sementes, enraizamento de estacas e anatomia da rizogênese em
Eugenia dysenterica DC. UFV, Viçosa.
Duarte E. F., Naves R. V., Borges J. D. & Guimarães N. N. R. (2006)
Germination and vigor of cagaita (Eugenia dysenterica Mart. ex
DC.) seeds as affected by seed size and harvest type. Pesquisa
Agropecuária Tropical 36 (3): 173–179.
Duke J. A. (1965) Keys for the identification of seedlings of
some prominent woody species in eight forest types in
Puerto Rico. Annals of the Missouri Botanic Garden 52 (3): 314–
350.
Duke J. A. (1969) On tropical tree seedlings. Annals of the Missouri
Botanic Garden 56 (2): 125–161.
Faria L. C. P., Carrara L. A., Amaral F. Q., Vasconcelos M. F., Diniz
M. G., Encarnação C. D., Hoffmann D., Gomes H. B., Lopes
L. E. & Rodrigues M. (2009) The birds of Fazenda Brejão: a
conservation priority area of Cerrado in northwestern Minas
Gerais, Brazil. Biota Neotropica 9 (3): 223–240.
Farias-Neto A. L., Fonseca C. E. L., Gomide C. C. C. & Silva J. A.
(1991) Storage of seeds of cagaita (Eugenia dysenterica DC).
Revista Brasileira De Fruticultura 13 (2): 55–62.
Ferreira M. B. (1973) Frutos comestíveis nativos do Distrito Federal.
Fundação Zoobotânica do DF, Brasília.
Finch-Savage W. E. & Leubner-Metzger G. L. (2006) Seed dor-
mancy and the control of germination. New Phytologist 171 (3):
501–523.
Franco A. C. (2004) Estratégias funcionais de plantas lenhosas das
savanas do Brasil Central. In: Cabrera H. M. (ed.). Fisiologia
Ecológica en Plantas. EUV, Valparaíso, pp. 173–188.
Gates B. (1982) Banisteriopsis and Diplopterys (Malpighiaceae).
Flora Neotropica Monograph, n 30. New York Botanical Garden,
New York.
Gross K. L., Maruca D. & Pregitizer K. S. (1992) Seedling growth
and root morphology of plants with different life-stories. New
Phytologist 120: 535–542.
Hoffman W. A. (1998) Post-burn reproduction of woody plants
in a neotropical savanna: the relative importance of sexual
and vegetative reproduction. Journal of Applied Ecology 35:
422–433.
Plant Species Biology 28, 130–137
137
Linkies A., Graeber K., Knight C. & Leubner-Metzger G. (2010)
The evolution of seeds. New Phytologist 186: 817–831.
Moreira A. G. & Klink C. A. (2000) Biomass allocation and growth
of tree seedlings form two contrasting Brazilian savannas.
Ecotropicos 13: 43–51.
Naves R. V., Borges J. D., Rocha M. R., Chaves L. J. & Vidal V. L.
(1992) Sprouting of seedlings of (Eugenia dysenterica) in glass-
house. Revista Brasileira De Fruticultura 14 (2): 37–40.
Oga F. M., Fonseca C. E. L. & Silva J. A. (1992) Influence of depth
and light in the germination of seeds of cagaita (Eugenia
dysenterica). Revista Do Instituto Florestal 4 (2): 634–639.
Palhares D. (2003) Pharmacognosy of the leaves of Eugenia dys-
enterica. Lecta-USF 21 (1–2): 29–36.
Palhares D., Paula J. E. & Silveira C. E. S. (2006) Morphology of
stem and subterranean system of Brosimum gaudichaudii
(Moraceae). Acta Botanica Hungarica 48: 89–101.
Palhares D. & Silveira C. E. S. (2007) Morphology of young plants
of Brosimum gaudichaudii Tréc. (Moraceae) cultivated in alter-
native conditions. Brazilian Journal of Medicinal Plants 9 (1):
93–96.
Pina G. O., Borghetti F., Silveira C. E. S. & Pereira L. A. R. (2009)
Effects of Eugenia dysenterica leaf extracts on sesame and
radish growth. Allelopathy Journal 23: 313–322.
Proença C. E. B. & Gibbs P. E. (1994) Reproductive biology of
eight sympatric Myrtaceae from Central Brazil. New Phytolo-
gist 126 (2): 343–354.
Rizzini C. T. (1970) Tegumentar effect on the germination of
Eugenia dysenterica. Revista Brasileira de Biologia 30 (3): 381–342.
Salomao A. & Allem A. (2001) Polyembryony in angiospermous
trees of the Brazilian Cerrado and Caatinga vegetation. Acta
Botanica Brasilica 15 (3): 369–378.
Sano S. M., Fonseca C. E. L., Ribeiro J. F., Oga F. M. & Luiz A. J.
B. (1995) Leafing, flowering, fruiting and initial growth of
cagaiteira in Planaltina-DF. Pesquisa Agropecuária Brasileira
30 (1): 5–14.
Silva R. S. M., Chaves L. J. & Naves R. V. (2001) Characterization
of cagaita fruits and tree in Southeastern region of Goiás State,
Brazil. Revista Brasileira De Fruticultura 23 (2): 330–334.
Souto L. S. & Oliveira D. M. T. (2005) Morphology, anatomy and
development of Byrsonima intermedia A. Juss. (Malpighi-
aceae). Brazilian Journal of Botany 28 (4): 697–712.
Souto L. S. & Oliveira D. M. T. (2008) Morphology, anatomy and
ontogeny of the seeds of Banisteriopsis C.B. Robinson e Diplop-
terys A. Juss. (Malpighiaceae). Acta Botanica Brasilica 22 (3):
733–740.
Souza E. R., Naves R. V., Borges J. D., Vera R., Fernandes E. P.,
Silva L. B. & Trindade M. G. (2008) Cagaiteira (Eugenia dysen-
terica DC.) phenology in Goiás State. Revista Brasileira De Fru-
ticultura 30 (4): 1009–1014.
Zaidan L. B. P. & Carrera R. C. (2008) Seed germination in
Cerrado species. Brazilian. Journal of Plant Physiology 20 (3):
167–181.
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