RELATIONSHIPS AMONG TIME, AND FLOWERING FREQUENCY

OF SOME FABACEAE TREE SPECIES

BRAMANTYO TRI ADI NUGROHO

BALI BOTANIC GARDEN, INDONESIAN INSTITUTE OF SCIENCES (LIPI)
Desa Candikuning, Kec. Baturiti, Kab. Tabanan – Bali
E-mail. bram001@lipi.go.id / bram_shearer@yahoo.com
Telp./fax. 0368 – 21273/ 22051 / 081310163035

ABSTRACT

Long term data on flowering phenology can provide insights into the temporal and
quantitative variation of resources availability for pollinators and in conjuction with weather data, and
several studies have shown that rainfall patterns significantly affect plant phenology. Flowering
patterns are defined by the timing, duration, and frequency of flowering. The objective of the present
study was to focus is on the timing and frequency of Fabaceae species flowering patterns and the
influence of each parameter on the other one. The study was conducted at Bali Botanic Garden for 5
years (2002-2006), encompassing 18 species and 12 genus of Fabaceae family. The highest Fabaceae
tree species that flowering was in June, October and December (77.8%), and the lowest in May and
August (50%). When flowering time of each species categorized according to the seasons, more than a
half of the observed trees were found in the both wet season, 12 species of 18. The species that
flowering episodically and supra-annually were more abundant (66.7 and 16.7%, respectively) than
those flowering continually and annually (11.1 and 5.6%, respectively). Flowering time and frequency
were significantly associated (G = 16.967, P < 0.01, df = 6). All of continually flowering species
flowered in both wet season. Continually flowering species were 1.2 more likely to flower in both wet
seasons than episodically flowering species.
Key words : Fabaceae, flowering, frequency, timing.
 INTRODUCTION

Plant phenology, and in particular flowering time, has been the focus of

many studies ever since the emergence of botany as a science (Rathcke and Lacey,

1985; Dahlgren et al. 2007). Timing of seasonal plant development can affect biotic

interaction and plant fitness (Dahlgren et al. 2007). Long term data on flowering

phenology can provide insights into the temporal and quantitative variation of

resources availability for pollinators and in conjuction with weather data, and several

studies have shown that rainfall patterns significantly affect plant phenology (Inouye

et al. 2003). Flowering patterns are defined by the timing, duration, and frequency of

flowering. Plants, particularly in the tropics, vary enormously with respect to these

main variables of flowering (Bawa et al. 2003). The extensive literature on flowering

phenology of tropical plants is largely focused on the timing or seasonality of

flowering at the community level (Opler et al. 1980; Wright and Calderon, 1995 Cit

Bawa et al. 2003).

The objective of the present study was to focus is on the timing and

frequency of Fabaceae specie flowering patterns and the influence of each parameter

on the other one. The research was done at Bali Botanic Garden. The focus of timing

and frequency of flowering base on the predictions from various hypotheses about the

evolution of flowering patterns, but have rarely been explicity tested in a

comprehensive manner. The Fabaceae or Leguminosae are a large and economically

important family of flowering plants. It is the third largest family of flowering plants

(after Orchidaceae and Asterraceae) with 730 genera and over 19.400 species,

according to the Royal Botanical Gardens. The largest genera are Astragalus (more

than 2.000 species), Acacia (more than 900 species), Indigofera (700 species),

Crotalaria (600 species), and Mimosa (500 species).

 Several predictions can be made about the timing and frequency of

flowering. Janzen (1967) suggested that tree species in dry tropical forest should

flower in dry season because the wet season is the major period for vegetative growth

for these species. Reproduction in the dry season allows temporal separation of

reproductive activity and vegettaive growth (Bawa et al. 2003). With respect to

frequency, in order to maximize lifetime reproductive output, species that flower

several times a year should be more common than species that flower annually (Bawa

et al. 2003).

In summary, our predictions are as follows : (1) Most tree species of

Fabaceae should flower during the period with the lowest rainfall (2) Species that

flower several times a year should be most abundant, and supra-annual species least

abundant.

MATERIALS AND METHODS

The study was conducted at Bali Botanic Garden for 5 years (2002-2006),

encompassing 18 species and 12 genus of Fabaceae family. This garden covers 154

ha, located in provinve of Bali, with an annual rainfall of 2000 mm. Flowering time,

and frequency were noted at weekly intervals for Fabaceae tree species.

Many trees species of Fabaceae at Bali Botanic Garden for several months

or several times a year, and the month of flowering of these species varied within a

range from year to year. Bawa et al (2003) used the method with pool the data over

the years. Thus, the month of flowering for these species was obtained from data

pooled over 5 years (2002-2006). Flowering time for each species was then

categorized into four level primarily on the basis of seasonality, in particular,

flowering in the “first wet” (Feb-Apr), “second wet” (Nov-Jan) and “dry” is the rest
of the month. Wet and dry month follow the differentiation by Wisnubroto et al

(1986), the month with more than 100 mm is wet month and less than 60 mm is dry

month. “First wet” consists of species that flower during the first wet season but do

not flower during the second wet season. “Second wet” consists of those species that

flower in the second wet season, but not in the first. “Both wet” consists of those

species that flower in the first and the second wet season or from the first to the

second wet season during a year. “Dry” consists species that flower only in dry season

during a year (Bawa et al, 2003).

Classification of flowering frequency follows the differentiation by Bawa et

al (2003). It was classified into four levels base on flowering frequency per year :

continual species that flower more or less continually during a year, episodic species

that flower more than once a year, annual species that flower once a year, and supra-

annual species that flower less frequently than once a year. Episodically flowering

species flower from two to six times a year (sub-annual flowering frequency). The

observations most likely reflect the minimum number of episodes within a year.

The relationship between flowering time and frequency categorical variables

was tested using two-way contigency table analyses. Bawa et al (2003) used the G

value (Likelihood ratio chi square ) to identify the categories contributing significanly

with examined adjusted standardized residuals.

RESULTS AND DISCUSSIONS

Description of flowering characters – Flowering time - The number of

flowering Fabaceae species are very varied among months (Fig 1). The highest

Fabaceae tree species that flowering was in June, October and December (77.8%),
and the lowest in May and August (50%). When the months were grouped into four

periods (February-April, May-July, August-October, November-January), the most

that flowering during May-July, and the fewest was August-October (Fig. 2).

When flowering time of each species categorized according to the seasons,

more than a half of the observed trees were found in the both wet season, 12 species

of 18. In the first wet season was no one of the species flowered. Thus, the proportion

of species flowering in each season decreased in order of both wet season (66.7%) >

dry season (22.2%) > second wet season (11.1%) > first wet season (0%). Bawa et al

(2003) showed because flowering time is based on data pooled by month for several

years, and the month of flowering can vary each year , most species belonging the

first wet, second wet and both wet categories flowered in dry season as well. In this

study, 100% of the species that flowering in the both wet season was also flowered in

dry season during the 5 years.

Fig 1. Number of flowering Fabaceae tree species during each month of the year. Flowering time of a species
varied within a range, and the month of flowering was pooled over 5 years (2002-2006)
Fig 2. Flowering Fabaceae tree species, grouped into four periods.

The hypothesis of most species flowering in the dry season was supported by

the data, even most of species was also flowering in the other months with high

rainfall (Fig.1). The highest of flowering species when the months were pooled into

four groups was May-July (Fig.2), when the lowest rainfall. When flowering time was

categorized according the season, the most abundant of flowering species was in the

both wet season. In this study, 100% of those species were also flowered in dry season

during five years, because flowering time is based on data pooled by month for five

years and the month of flowering can vary each year. Thus, in fact, the most abundant

of flowering species was in dry season.

The most species flowered in dry season because the pollinators are more

abundant in the dry season than wet season, and most of Fabaceae species was

strongly assosiated with medium or large bees (Bawa et al. 2003; Kang and Bawa,

2003). An other reason, in the wet season is the major period for vegetative growth for

these species. Many species is also flowered in dry season because in this season leaf

of these species that fall to reduce stress (Borchert, 1983 Cit Bawa et al 2003).
Observed phenological patterns range from dought deciduous to evergreen trees

which exchange leaves and flowers during the dry season. Within four weeks after the

last major rainfall, the upper soil layers dry out and tree water balance becomes

negative in all trees, as indicated by declining stem water potential. In many trees,

elimination of water loss by leaf shedding in combination with with absorptin of

water from the subsoil (Borchert, 1998). Variation in phenology of Erythrina

poeppigiana showed that this species flower in dry season at the same time with leaf

shedding (Borchert, 1998).

Flowering frequency - The episodically and supra-annually flowering

species were more abundant (66.7 and 16.7%, respectively) than those flowering

continually and annually (11.1 and 5.6%, respectively). In episodically flowering

Fabaceae species, episodes varied from two to six times per year, 50 % of species

with episodic mode flowered twice times a year (Fig. 3).

Fig 3. Proportion of episodically flowering species with different flowering frequencies per year. N = 12

The hypothesis that species that flower several times a year should be more
abundant than species that flower annually was not fully supported by the data

because supra-annually species were more abundant than continually and annually

species. Some studies have shown that supra-annually usualy the least abundant.

Bawa et al (2003) described that flowering once a year is likely to confer a

disadvantage because unpredictability of flowering will inhibit pollinator fidelity and

specialization. There is probably the reason to explain the supra-annually flowering

species were more abundant than continually and annually flowering species. Most

pollen vector of Leguminosae was medium or large bee, and pollen vector for supra-

annual species was also medium or large bee and the pollen vector of supra-annually

species was more abundant than continually species (Kang and Bawa, 2003).

Many plant species in the tropics display episodic flowering, little is known

about its adaptive (Bawa et al 2003). Newstrom et al (1994) Cit Bawa et al (2003)

reported that episodic flowering is more frequent than annual flowering species.

Episodic flowering, may be constrained by fruit development and maturation and

optimal timing for seed dispersal. If seeds and fruits mature over a long period of

time, repeated flowering would not be posible. Similarly, if seed dispersal is seasonal,

as is dispersal by wind during the dry season, there would be slection for annual

flowering (Bawa et al 2003). Continually flowering species is rare is not surprising.

Continual flowering, which necesarily accompanies continual fruiting, requires

special condition to ensure resource acquisition. Indeed, these species tend to occur in

patchy, sunlit habitats (Stiles, 1978; Opler et al 1980 Cit Bawa et al 2003).

Relationship between flowering characters – Flowering time and frequency

(Table 1) were significantly associated (G = 16.967, P < 0.01, df = 6). All of

continually flowering species flowered in both wet season. Continually flowering

species were 1.2 more likely to flower in both wet seasons than episodically flowering
species. In contrast, no one of the annual and supra-annual flowering species flowered

in both wet season. Supra-annually flowering species were 12 times more likely to

flower in the dry season than episodically flowering species (Table 1).

Table 1. Frequency of Fabaceae tree species at Bali Botanic Garden, in a two way contingency table of flowering
time and frequency.
Flowering time
First Scond Both Dry
Trait wet wet wet G
Flowering frequency

Conitinual 0 0- 2+ 0-
Episodic 0 1- 10+ 1-
Annual 0 1+ 0- 0-
Supra-annual 0 0- 0- 3+ 16.967**
Note : +/- signs indicate that observed values were greater or less than the expected values at alpha = 0.05 level.
** P < 0.01.

Relationship between flowering time and frequency – The prediction that

supra-annual species should flower in dry season. Several authors have shown that

supra annual species flower massively mainly in the dry season (Frankie et al. 1974;

Opler et al. 1980; Ashton et al. 1988 Cit Bawa et al. 2003). Supra annual flowering

may have evolved in response to selection to attract pollinators or to satiate seed

predators. Supra annual flowering in Bornean tropical forest has also been interpreted

as a mechanism to increase mating opportunities by utilizing diverse general vectors

and to reduce seed predation (Bawa et al. 2003).

CONCLUSIONS
We have shown that there is association among flowering character,

relationship between flowering time and frequency. The association was showed that

the G value, and all the supra annually flowering species flowered in dry season. The

second general point is that approximately 66.7% of all Fabaceae species at Bali

Botanic Garden flower episodically, and 50% of them flower two times a year. The
most species flower in dry season because pollinators are more abundant in the dry

season than wet season, and most of Fabaceae species was strongly assosiated with

medium or large bees.

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