Reproductive phenology of Carapa guianensis Aubl. (Meliaceae) in two forest areas of the Central Amazon

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International Journal of Environment, Agriculture and Biotechnology (IJEAB) Vol-3, Issue-3, May-June- 2018 ISSN: 2456-1878 Page | 714
Reproductive phenology of Carapa guianensis
Aubl. (Meliaceae) in two forest areas of the
Central Amazon
Antenor Barbosa1; Antonio Moçambite1; Patrícia Morellato2 and Cláudia Blair e Matos1
1Instituto Nacional de Pesquisas da Amazônia INPA/COTEI Manaus, AM.
2Universidade Estadual Paulista-UNESP - Departamento de Botânica, Rio Claro, SP. [email protected]
Abstract This article presents the phenological study of
Carapa guianensis Aubl species from 1974 to 2000, in
ADFR and TFES forests stations research in Central
Amazon, Brazil. The objective was to analyze and
compare the phenological pattern (flowering and
fructification) and the influence of the climatic factors.
The flowering in the TFES started in a higher
precipitation season; meanwhile at ADFR it was
irregular. The fruiting in both areas occurred more
frequently rainiest season, but in the ADFR the mature
fruits were mo re irregular. The frequency of occurrence
was annual from “flower bud” to “immature fruit”
phenophases in TFES, but was over-annual only in
“mature fruits”. But in ADFR, was annual from “flower
bud” to “anthesis” and was over-annual in immature
fruit” and “mature fruit”, both with irregular pattern and
duration from intermediate to p rolonged. The duration of
the floral bud phenophase and anthesis was similar in the
two areas; however, ‘immature fruits” in the TFES, in
general, was higher than in the ADFR. But “mature
fruits” were higher in ADFR. The phenophases did not
occurred at same time in all trees studied, possibly due
the influence of the intraspecific genetic variability in
interaction with the environment.
Keywords crabwood, flowering, fruiting, raining
The long-term phenological study began in 1963
in the Adolph Ducke Forest Reserve (ADFR) and in 1974
at the Tropical Forest E xperiment Station (TFES), both
located in the region of Manaus, Amazonas, Brazil [1, 2].
In these areas, there is a major investigation of
phenological events of about 120 species of tropical
forests to generate subsidies to the management and
reforestation plans in the Amazon forest. This research
has generated several publications on the phenolo gy of
trees in the Central Amazon, which analyze the data
collected in the ADFR. The former studied the phenology
of Amazonian forest species occurred in a period of seven
years (1962-1968) and the latter analyzed the phenology
of forest species in upland tropical rainforests in the
Central Amazon [1, 2, 3, 4, 5, 6, 7, 8].
In what concer ns to the population of a single
species or family, studies present results on the phenology
of Aniba rosaeodora Ducke (Lauraceae) for a period of
eleven years (1968-1978) [3]. T he phenology of
Copaifera multijuga Aubl (Fabaceae) was evaluated in
seven years (February 1979 to December 1985), and there
in relation to weather elements [4]. Also, was analyzed
the phenological behavior of Diplotropis purpurea Rich.
(Fabaceae) in six years (1980-1985 ) [5]. Among the
studies on families, two must be highlighted: the
phenology of five species of Lecyt hidaceae over an
eleven-years period (1978-1988) [6] and the p henology
analysis of five species of Sapotaceae in one twenty-one-
year period (1970-1990) [7].
The species Carapa guianensis Aubl. (Meliaceae),
commonly known as crabwood or andiroba in Br azil, was
selected for the analysis of phenological behavior, due to
its economic, social and ecological importance, also
abundant in the Amazon region. The trees of andiroba can
reach up to thirty meters high with a cylindrical trunk,
straight and buttresses at the base [8,9]. It is a species of
multiple uses, having a high-quality wood, which can be
used in carpentry, construction, shipbuilding, boards and
plywood, furniture, beams, interior works, pencils, masts
and others. Another extraor dinary use is the oil extracted
from its seeds, which is currently one of the most
important products in the regional market. Andiroba oil is
a clear and transparent liquid, that at temperatures b elow
25°C it solidifies as vaseline [9,11, 12].
The oil can be used in the manufacture of soaps,
candles, in the composition of cosmetics and in different
medicines because andiroba oil is a rich source of
essential fatty acids, including oleic, palmitic, myristic
and linoleic acids, and contains no fatty components such
as triterpenes, tannins, a nd alkaloids. The bitter taste of
International Journal of Environment, Agriculture and Biotechnology (IJEAB) Vol-3, Issue-3, May-June- 2018 ISSN: 2456-1878 Page | 715
the oil is attributed to a group of terpene chemicals called
meliacins, which are very similar to the bitter antimalarial
chemicals. Recently, one of these meliacins, called
gedunin, was documented to have pest control properties
and antimalarial effects equal to that of quinine. A
chemical analysis of andiroba oil identified the anti-
inflammatory named andirobina, which has healing and
insect repelling properties that are attributed to the
presence of limonoids. The interest in using andiroba oil
in cosmetics has increased significantly, especially after
the patenting of a cream by Yves Rocher, from France,
that has moisturizing and anticellulite p roperties based on
this oil. [13].
The phenological study of this species is essential,
as it enables the determination of the regularity and
predictability in the supply of this natural resource, which
allows more rational use in the Amazon.
The phenological patterns would be most affected
by the intrinsic characteristics (ge netic, physiological and
reproductive) of the species and by ecological factors
(pollination, predation, competition) and not only by
climate variables [7]. Researchers also report the
influence of climatic elements (precipitation, solar
radiation, evaporation, relative humidity) basis o n
phenological studies on Copaifera multijuga and five
Sapotaceae species [4, 7].
Phenology studies were installed in ADFR in 1963
and in TFES in 1970. Since then, phenophases of
flowering, fruiting and leaf change have been studied.
However, the ADFR is no longer surrounded by native
forests, because all the perimeter has been deforested by
the urban e xpansion in Manaus city. While the EEST is
43 km away from the nearest town and surrounded by
native forest. Consequently, the climatic conditions and
the interaction with other biotic components, especially
pollinators, predators and dispersers, are under different
conditions and, therefore, may influence the patterns of
occurrence of the phenophases studied in this work.
This study aimed to determine the patterns of
flowering (flower bud and anthesis) and fruiting (mature
and immature), of C. guianensis and compare the
phenological events in order to determine whether this
species has similar phenological behavior in two distinct
areas of upland forest (ADFR and TFES) and if it
responds to the climatic factors over time between the
years from 1974 to 2000. It is important to determine the
pattern of flowering and fructification of phenophases to
characterize the ecological group of forest succession
(climax) to which these species belong [14]. The
knowledge of the phenological pattern allows more
specific studies on the reproduction of the species and
also to provide basic information to support the planning
of silvicultural projects for species plantations, timber and
oil prod uction and for the recovery of degraded areas,
since there are good growth results i n experimental
plantations [15,16 ,17]. Therefore, the hypothesis of the
study is that the climatic change s affect the phenophases
of the Amazonian species and impair the prod uction of
fruits, thus reducing the supply of seeds for trees’
reproduction and reducing the source of food for animals,
changing the forest’s ecological balance.
The studies were conducted in the ADFR, located
26 km north of Manaus, on AM-010 Road, measuring
10,072 ha in an upland rainforest at 59o52’40” to
59o52’00” west longitude and 03o00’00” to 03o08’00”
south latitude [18] and in the TFES, located at
approximately 45 km north of Manaus, on BR-174 Road,
measuring 21,000 ha, at 2o37’ to 2o38’ south latitude and
60o09’ to 60o11’ west longitude [19].
According to the Köppen classification system,
local climate is designated Afi: A - tropical climate with
virtually no winter, the average temperature for the
coldest month is never lower than 18oC; f - rai ns
throughout the year; i - indicating isotherm, that is, the
annual average temperature fluctuations do not reach 5oC;
there is no winter or sum mer [19]. Climatological data
used in this study were provided b y the Coordination of
Research on Environmental Sciences of the National
Institute of Amazonian Research (INPA) and collected at
the climatological station of the ADFR for the two
experimental areas, which is located approximately 30 km
away from the TFES.
The Figure 1 shows annual rainfall and minimum,
average and maximum temperatures from in twenty-seven
years data (1974-2000).
The driest month was August with 101 mm and
the month with the highest average precipitation was
April with 304.34 mm. T he average monthly temperature
ranged from 25.5°C to 26.7oC. T he average of minimum
temperatures that predominated were around 22°C.
Maximum temperatures ranged from 31.3oC to 33.4oC at
the end of the dry season, whereas the lo west values were
observed in the rainy season 22,1oC.
The frequent rainfall which extend from
November to May, called rainy season, reached monthly
averages over 263 mm and lower average temperatures o f
25,8o C. There is also a less humid period between June
and October, with less constant rainfall, but no water
deficit. In this period, there was an average rainfall of 121
mm per month and higher temperatures of 32,5 o C,
considered as the dry season.
The predominant vegetation in the region was
classified as a tropical upland rainforest, characterized by
a great diversity of tree, shrub and herbaceous species
International Journal of Environment, Agriculture and Biotechnology (IJEAB) Vol-3, Issue-3, May-June- 2018 ISSN: 2456-1878 Page | 716
The forest that covers the areas studied in t his
work is part of the Amazon Moist Forest [22] which is
always green, as the trees never lose all the foliage, at the
same time, and has a large number of tree species that are
usually divided into three distinct strata.
The upper or dominant stratum is formed b y large
trees with DBH (diameter at breast height) greater than 1
m and height sometimes reaching 45 m or more, as
happens with Cedrelinga catenaeformis Ducke
(Mimosoideae) and Dinizia excelsa Ducke (Fabaceae) .
The intermediate stratum (vegetation layer) is composed
of smaller trees, whose DBH may exceed 1 m, but their
height is usually below 45 m, as happens with
Enterolobium schomburgkii Bth (Fabaceae), Aniba duckei
Kostermans (Lauraceae), and palm trees s uch as Euterpe
oleracea Mart. (Arecaceae) and Mauritia aculeata H.B.K.
(Arecaceae). The lower stratum consists of species that
develop in heavy shade conditions, such as Geonoma
deversa (Poit) Kunth. (Arecaceae), Manicaria saccifera
Gaertn (Arecaceae) and other shrubs and herbaceous
plants [23].
The trees of the phenological study were
previously selected in the forest according to their habitat,
height, DBH and stem form [1]. Five C. guianensis
individuals were sampled from the ADFR and five from
the TFES. Subjects were observed with the aid of
binoculars to record the phenological phases. Monthly
observations were carried out in this study. The following
phenophases were analyzed: flowering and fruiti ng. The
flowering was divided in “flo wer buds” (appearance) and
“anthesis” (earl y flowering). The Fruiting, divided into
“immature fruits” (appearing new fruits) and “mature
fruits” (presence of ripe fruits). The analysis was done
from data collected monthly to verify the frequency of
events, from 1974 to 2000 [2].
Phenological patterns are described according to
“frequency” - number of cycles with and without
phenophases per year, “reg ularity” - variation in the time
of occurrence and, "duration of cycles or phases" - time in
months that an individual remains in a phase or cycle
The repetitive occurrence of phenological events
in the year is called “annual frequency”. According to the
annual frequency of flowering and fruiting, the species
are classified as: “sub-annual” (more than one event per
year), “annual” (one event ea ch year) and “over-annual”
(events at intervals of two years or more) [24].
The phenological data of ADFR and TFES were
stored in DBASE III software and analyzed in
FENOLOG, which is a software developed at the
Coordination of Research on Tropical Forestry of INPA.
The relationships between phenological data and climate
variables was calculated by the non-parametric analysis of
Spearman's correlation coefficient, considering the
monthly average values of climate variables [25].
3.1 Occurrence and pattern of phenophases
3.1.1 “Flower bud” phenophase
The “flower bud” phenophase of C. guianensis in
TFES presented annual frequency, normally occurring at
the beginning of the rainy season and positive correlation
with the minimum temperature (rs = 0.11, p <0.05) and
day length (rs = 0.24, p <0.01).
The greatest number of trees (3-4) with “flower
bud” per month o ccurred in the years 1975 (Jan), 1976
(Dec), 1977 and 1978 (Nov), 1979 (Set), 1980 (Oct),
1983 (Jan), 1984 end 1986 (Oct), 1987 (Dec) and 1988
(Nov). In the years 1981, 1985, 1989, 1990 and 1991,
there were no trees with flower buds. Only in 1992 two
trees produced bud flo wer, but only one tree in the years
1982, 1992 until 1999 (Fig 2A).
The peaks of occurrence (three or more trees per
year) were registered in 1975 (Jan and No v), 1976 (Nov
and Dec), 1977 and 1978 (Nov), 1979 (Feb, Sep and Oct),
1980 (Oct and Dec), 1983 (Jan), 1984 and 1986 (Oct),
1987 (Dec), and 1988(Nov) (Fig 2A).
The “flower bud” phenophase of C. guianensis in
ADRF presented annual frequency occurring at the
beginning of the rainy season, and positive correlation
with the minimum temperature (rs=0,12; p<0,05) and day
length (rs=0,21; p<0,01).
The greatest number of trees (3-5) with “flower
bud” per month occurred in the years 1976 (Jan, Nov and
Dec), 1 977 (Nov and Dec), 1979 (Feb, Ago, Sep and
Oct), 1989 (Nov), 1980 (Oct) But without flower bud,
were observed in 1975, 1978, 1 980, 1986, 1988, 1990,
1991, 1992, 1994, 1997 e 1998. Only in 1985 (Jun), 1995
(Nov and Dec) and 1999 (Sep) two trees produced bud
flower, but only one tree in the years 1974 (Jul),1976
(Jan), 1979 (Aug, Sep and Oct), 1981 and 1982 (Nov),
1983 (Oct and Nov), 1984 (Feb, Oct, Nov and Dec), 198 7
(Jul), 1993 (Aug), 1996 (Oct), 1999 (Oct and Nov) and
2000 (Nov) (Fig 2B).
The peaks of occurrence were registered in 1976
(Jan, Nov and Dec), 1977 (Nov and Dec), 1979 Feb, Aug,
Sep an Oct), 1984 (Feb, Oct, Nov and Dec), 1989 (Nov),
1995 (Nov and Dec), 1999 (Sep, Oct and Nov) (Fig 2B).
3.1.2 - Anthesis phenophase
The Anthesis of C. guian ensis in TFES, had the
tendency of usually starting during the higher
precipitation season and presented a positive correlation
with minimum temperature (rs = 0.14; p < 0.01).
The greatest number of trees (3-5) per month
occurred the anthesis in the years 1975 (Jan and Feb),
1976 (Dec), 1977 (Nov and Dec), 1978 (Nov), 1979 (Sep