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ORIGINAL ARTICLE

International Journal of Environment, 1(1): 2127 (2011)

Vegetative and Reproductive Phenology, and Litterfall Production of

Rhizophora stylosa in Okinawa Island, Japan
Sahadev Sharma, Md. Kamruzzaman, ATM Rafiqul Hoque,
Kangkuso Analuddin and Akio Hagihara
Abstract

ISSN: 2186-6872 (print)

ISSN: 2186-0009 (online)
http://www.BENJapan.org/IJE

ARTICLE HISTORY
Received: 14 April 2011
Revised: 20 May 2011
Accepted: 28 May 2011

Mangrove forests in Japan are at their northern limits, and are

distributed from the Yaeyama Islands in the subtropical zone to the
southern part of Kyushu Island in the warm-temperate zone, and
therefore they are poor in species. Mangroves growing in subtropical
region have very few knowledge regarding phenology. This species is
growing in its northern limit and ambient climatic conditions, so
therefore phenology and litterfall studies are needed to know its
adaptation strategies and growth. In this study, Phenological traits of a
subtropical Rhizophoraceae mangrove, Rhizophora stylosa Griff., are
measured during the period of April 2008 to March 2010. The litterfall
traps were emptied monthly and the litterfall was sorted into leaves,
stipules, branches, flower buds, fruits, fruits expanded and hypocotyls
were counted monthly. Litterfalls showed a seasonal pattern: the highest
peak occurred during summer and the lowest during winter. Mean
annual total litterfalls of the 1 st year and 2nd year were respectively
estimated as 931.716.6 and 1321.931.2 (SE) g m -2 yr-1. Leaf litterfall
comprised 65.2 (1st year) and 49.7% (2nd year) of the total litterfall.
Conversion of fertilized flowers into fruits was greater than half (75
90%), though transition of fruits to propagules was less successful (1.3
3.3%). Therefore, we can conclude that the survival rate of reproductive
units was low, with very few of flowers converted into propagules (1.2
2.5%). The reproductive cycle of Rhizophora stylosa was seasonal. In
general, the species presented flowering peaks from May to September
when temperature was high.
Keywords: Northern limit, Resource limitation, Reproductive cycle, Survival rate.

Published online: 29 July 2011

2011, International Journal of Environment. All rights reserved.

AUTHORS INFO

I.

Sahadev Sharma*
e-mail: mangrove_coral@yahoo.co.in
Graduate School of Engineering and Science
University of the Ryukyus, Okinaw, Japan.

INTRODUCTION

The floral biology of mangroves in general has

received little attention, and highlights the
need for quantitative and experimental
information for understanding the ecology and
genetics of a group of species that are of
considerable environmental importance. The
reproductive biology of mangroves has often
been regarded with interest owing to the
unusual breeding mechanisms, including
vivipary, exhibited by many of these plants [1].
In fact, the focus of research on mangrove
reproductive biology has almost exclusively
been on the fruit dispersal stage [2]. Little is
known about the reproductive processes and
success rate of fruit and propagule production
in mangrove ecosystems.
Mangroves are unique trees that are found
almost exclusively in the tropics and
subtropics. Mangroves frequently show a high
within-region diversity of structural patterns,
and an equally high diversity of functional
roles [3, 4]. Based on knowledge of the study
sites, high values for height, biomass, and

Md. Kamruzzaman
e-mail: zaman_dulu@yahoo.com
Graduate School of Engineering and Science
University of the Ryukyus, Okinawa, Japan.
ATM Rafiqul Hoque
e-mail: atmrafiqul@gmail.com
Institute of Forestry and Environmental Sciences
Chittagong University, Chittagong, Bangladesh.
Kangkuso Analuddin
e-mail: udin_zanza@yahoo.com
The Wallance Research Center, Heluoleo
University, Kendari, Indonesia.
Akio Hagihara
e-mail: amyh@sci.u-ryukyu.ac.jp
Faculty of Science, University of the Ryukyus,
Okinawa, Japan.
* Corresponding author

21

-S Sharma et al.
litterfall
indicate
optimum
habitats,
particularly with respect to reduced salinity
regimes, more optimal climatic conditions, and
better site fertility [5]. Phenological data are
essential to know the tree ability to adapt
growth and propagation strategies to ambient
climatic conditions. This kind of knowledge is
most valuable if available across a broad
geographic scale. The litterfall is a main
component of net primary production [6],
reflects phenological events [7, 8] and is also
an important part of energy and nutrient
fluxes in mangrove ecosystems [9]. Rhizophora
are considered the most important of all
mangrove genera across the Pacific tropical
and subtropical region. However, little is
known about R. stylosa phenology in Okinawa
Island near the northern limit of mangrove
distribution; information is available for
Kandelia obovata and Bruguiera gymnorrhiza
[10, 11].
Many studies have documented mangrove
litterfall to assess mangrove productivity;
however, fewer studies have documented
mangrove phenology, particularly reproductive
phenology,
particularly
reproductive
phenology. Some studies has been done such
as, growth and reproduction in two species of
mangrove Avicennia marina and Aegiceras
corniculatum, measured over three years in
Jervis Bay, New South Wales, by demographic
censuses of leaf and floral components
supplemented by litterfall collections [8].
Knowledge
of
mangrove
phenology
is
important for understanding the mangrove
growth, productivity and life cycle as well as
plant - animal interaction with in the
ecosystem. As mangroves in Okinawa thrive in
a distinct seasonal climate, they have to cope
with
substantial
seasonal
changes
in
environmental factors. In general, R. stylosa
focused on this study shows a conspicuous
seasonality regarding phenophase peaks. The
aim of this study is to investigate the
vegetative and reproductive biology, and the
litterfall production of mangrove R. stylosa
growing in the subtropical region. The general
trends that regulate the vegetative and
reproductive production in the mangrove
system are discussed.
II. MATERIALS AND METHODS
A. Study site
The study was conducted in a mangrove
forest (26o11N and 127o40E) of Manko
Wetland, Okinawa Island, Japan (Fig. 1), from
April 2008 to March 2010. This wetland is an
important area for migratory birds and has

22

been registered as the RAMSAR site since

1999. The mangrove Kandelia obovata (S., L.)
Yong is the dominant species in the study site.
A few pure patches of Rhizophora stylosa
Griff., Bruguiera gymnorhiza (L.) Lamk. and
Excoecaria agallocha L. are also observed.
The warmth index [12] based on the data of
20002009 obtained from the Okinawa
Meteorological Observatory, Naha, Okinawa,
was 219.80C month, indicating that this area
belongs to the subtropical region. Monthly
mean minimum temperature is 160C in
January and monthly mean maximum
temperature is 29.40C in July during the
study period. Mean annual rainfall is 1417.5
mm yr-1 during the study period.

Urban area

Fig. 1. Location of the study area; the

hatched zone indicates the mangrove
area

B. Litter collection
Five plots (4 m x 4 m each) were established
for a Rhizophora stylosa pure stands. All
individuals in the subplots were counted and
numbered. Litterfall was measured using litter
traps. One litterfall trap with an opening of
0.19635 m2 was placed at 1 m above the soil
surface in each of 5 plots. The litterfall traps
were emptied monthly from April 2008 to
March 2010 and the litterfall was sorted into
leaves, stipules, branches, flower buds,
flowers, fruits, fruits extended (immature
hypocotyle with fruit part) and hypocotyles.
The sorted litterfalls were dried to a constant

Reproductive phenology and litterfall production of Rhizophora stylosa

mass at 80oC over a 48 h period, and then

weighed.
Table 1. Annual litterfall of the 1st year (Apr.2008
Mar.2009) and the 2nd year (Apr. 2009 Mar.2010).
Litterfall

1st year

2nd year

Leaf

610.059.54

657.2613.17

Stipule

104.331.76

111.062.24

Branch

68.524.31

107.104.96

8.820.29

12.340.37

Flower

31.961.21

43.271.42

Fruit

49.971.08

114.083.26

Fruit extended

15.560.85

29.441.68

Hypocotyle

46.512.80

247.3814.14

935.7316.68

1321.931.20

Flower bud

Total
Mean value (g

m-2

yr-1)

is shown with its SE.

III. RESULTS AND DISCUSSION

A. Vegetative phenology
As depicted in Fig. 2, leaf litterfall showed a
seasonal pattern: the highest peak occurred in
July and the lowest peak occurred during the
December. According to May, the leaf litterfall
of Avicennia marina (Forsk.) Vierh increased
with increasing air temperature [13]. In the
subtropical region, leaf fall and leaf production
are low during the winter season [1417]. At
lower latitudes, pattern of leaf fall and
production are less uniform [18]. In temperate
mangroves, leaf litterfall has been reported to
be unimodal and highest in summer [7], [13].
Although peak litterfall have been observed in
other mangrove wetlands during the rainy
season [19, 20]. Unimodal and bimodal peaks
in mangrove leaf production are known to be
associated with forests growing in higher and
lower intertidal areas respectively due to
fluctuations in ground water salinity [21].
Ochieng and Erftemeijer reported unimodal
leaf flush pattern in A. marina at Gazi bay
during wet season [22]. Saenger and Moverley
suggested that leaf production in mangroves
was closely related to changes in temperature,
with seasonal increase in temperature
associated with leaf initiation [23].
Stipule litterfall was the highest in July,
while the lowest during winter (Fig. 2). It is
obvious from Fig. 2 that new leaves were
formed or expanded at different rates
throughout the year, whereas shedding of old
leaves took place at more even rates.
Rhizophora mangle L., studied in subtropical

23

Florida by [14], is a continuously growing

species but shows a unimodal growth with a
very high rate of leaf replacement in the warm
and humid summer. Several studies in
Australia and Papua New Guinea have
reported marked seasonal periodicity in leaf
emergence for Rhizophora spp., with a peak
during the wet season [24, 25, 26]. Rates of
leaf emergence were significantly higher in the
wet season than the dry seasons, as observed
in Mekong delta, Vietnam [27]. Seasonality in
leaf emergence is less clear, as [24] did not
observe a seasonal periodicity in stipule
production based on the dry mass collected
month at sites on Hinchinbrook Island [28].
Although branch litterfall did not show any
clear seasonal pattern (Fig. 2), it showed the
highest peak during summer. According to
past studies, the highest branch litterfall of K.
obovata in Okinawa Island resulted from the
typhoon [29] and the branch litterfall of A.
marina depended on the storm [30]. Total
litterfall showed a seasonal trend with the
highest litterfall in summer and the lowest in
winter (Fig. 2). Mean annual total litterfalls of
the 1st year and the 2nd year were estimated as
935.716.7 and 1321.931.2 (SE) g m-2 yr-1
(Table 1), of which leaf litterfall comprised the
largest component, i.e. 65.2 and 49.7%
respectively in the 1st year and the 2nd year.
The mean annual litterfall recorded in this study
was within the range of values for Rhizophora
spp. in Australia and Vietnam [28, 31].
Table 2: Numbers of flowers, fruits, hypocotyles and
fruits extended during 1st year (Apr. 2008 Mar.
2009) and the 2nd year (Apr. 2009 Mar. 2010).
Litterfall

Flower

Fruit

Hypocotyle

Fruit
extended

1st year

422

383

2nd year

801

602

20

13

B. Reproductive phenology
Reproductive material was present in litter
samples all over the year. Flower bud,
comprising the bud primordia, immature bud
and mature bud, showed the highest peak
during summer and the lowest was in winter
(Fig. 3). Rhizophora stylosa has primordia and
flower buds with resting periods, about one
year each, a long period of around 2 years
being required for development of flower.
Rhizophora apiculata Blume needed 2 and half
years to develop flower [32]. Bud production
occurs throughout the year in Rhizophora
apiculata in Southern Thailand, although

-S Sharma et al.

common in tropical trees with comparatively

large fruits [32].

120
100

-2

-1

Leaf litterfall (g m month )

140

80
60
40
20
0
AM J J AS OND J FMAM J J AS OND J FM

30
25

-2

-1

Stipule litterfall (g m month )

35

20
15
10
5
0

AM J J AS OND J FMAM J J AS OND J FM

-1

Branch litterfall (g m month )

100

-2

80

60

40

20

0
400

AM J J AS OND J FMAM J J AS OND J FM

350

-2

-1

Total litterfall (g m month )

flowering occurs during the dry season [32].

Other
studies
have
reported
varying
reproductive cycles at different latitudes, e.g.,
five months in Darwin Harbour [33], six
months in southern Thailand [34] and one
year in southern-eastern Australia [35].
For Rhizophora magle, more or less
continuous
flower
production
with
a
superimposed seasonal trend has been
reported by several authors. Flower also
showed the highest peak during summer,
while the lowest was in winter (Fig. 3). Low
temperatures generally reduce the flower
production of R. mangle [14]. As with A.
marina, the differences in timing of flowering
among locations may be influenced by climatic
factors. In all species in Darwin Harbour, with
the exception of R. stylosa, peak flowering
occurs during the dry season through to the
start of the wet, consistent with the reported
by [25]. The dry conditions are likely to
facilitate pollinators and pollination.
Fruit, which comprises both immature and
mature fruits, also showed the highest
production during summer, while the lowest
was in winter (Fig. 3). It is reported in Florida
that fruit development extends over the cold
season and is therefore slower [14].
Hypocotyle and fruit extended found in only
from July to September (Fig. 3). In other
Rhizophoraceae
species,
propagule
development may last considerably longer [25,
27].
Rhizophora stylosa produced a large number
of buds, however, most buds failed to develop
into propagules. A large proportion of young
buds became mature buds, with most mature
buds becoming flowers. There was a sharp
decline in survival of reproductive units, with
very few of flowers fertilised (1.22.5%).
Conversion of fertilised flowers into fruits was
greater than half (7590%), though transition
of fruits to propagules was less successful
(1.33.3%) (Table 2). In Northern Australia and
Fiji, Rhizophora spp. has 13% of survival rate
for flower to propagule production [33, 36].
The rate of fertilisation reported in this study
was much lower than those estimated in R.
mangle (7%) and R. apiculata (13%) [37], but
similar to that for R. stylosa in north eastern
Australia (4%) [31].
Immature dropping of propagules may be
due to various reasons, such as insect damage,
non-availability of nutrients at the time of
development and tree size. This is also true
with several tropical/subtropical tree species
whose number of flowers is quite large as
compared to fruit set percentage [37]. Low
conversion of flowers to fruits is a feature

24

300
250
200
150
100
50
0

AM J J AS OND J FMAM J J AS OND J FM

2008
2009
2010
Month

Fig. 2. Seasonal trends in monthly mean litterfall.

Vertical bar stands for the standard error of the mean

Reproductive phenology and litterfall production of Rhizophora stylosa

-2

-1

Flower bud litterfall (g m month )

4
3
2
1
0

-2

-1

Flower litterfall (g m month )

16

AM J J AS ON D J FM AM J J AS ON D J FM

14
12
10
8
6
4
2
0
40

AM J J A S ON D J F MAM J J A S ON D J FM

-2

-1

Fruit litterfall (g m month )

35
30
25
20
15
10
5
0
AM J J A S ON D J F MAM J J A S ON D J FM

-1

Fruit extended litterfall (g m month )

25

-2

20

15

10

AM J J A S ON D J F MAM J J A S ON D J FM

-1

In summary, patterns in reproductive and

vegetative phenology are repeated each year in
R, stylosa. The seasonality and timing of these
patterns is likely to coincide with specific
climatic conditions, such as temperature. All
litterfalls showed a seasonal pattern: the
highest peak occurred during summer and the
lowest during winter. Mean annual total
litterfalls of the 1st year and 2nd year were
respectively estimated as 931.7 16.6 and
1321.9 31.2 (SE) g m-2 yr-1, of which the leaf
litterfall comprised 65.2 (1st year) and 49.7 %
(2nd year). The reproductive cycle of R. stylosa
was seasonal. Rhizophora stylosa requires a
long period of around 2 years for development
of flower. The survival rate of reproductive
units was low, with very few of flowers
converted
into
propagules
(1.22.5%).
Rhizophora stylosa hypocotyle is largest in the
family Rhizophoraceae. Consequently, the low
rate of fruit set recorded in this study may be
reflection not only of restricted pollination, but
also of resource limitation. The periodicity of
reproductive and vegetative phenology in
Manko Wetland mangroves is not consistent
for the other region as different factors
regimes within this large biogeographic region
are likely to result in intraspecific variations in
R. stylosa phenology.

150

-2

IV. CONCLUSION

Hypocotyl litterfall (g m month )

The low rate of fruit set recorded in this

study may be a reflection not only of restricted
pollination, but also of resource limitation. As
the propagules characteristic of R. stylosa are
likely to be metabolically expensive to produce,
trees may limit their output per reproductive
season, or abort the developing propagules
when energy reserves are restricted. Large
investments of resources to develop large
fruits are common to many mangroves [38].
The consequences of this phenomenon
appear to be that growth is determined by
maternal investment into fruits, as evidenced
by synchronized leaf and floral phenologies.
Thus, at high latitudes growth restricted not
only by a shorter growing season but also by
the resource demands of a large fruit that
takes longer to develop.
The study has thus shown that the vegetative
and reproductive phenology of R. stylosa at
Manko Wetland is clearly seasonal, evidenced
by peaks in the various phenophases. However,
vegetative peaks appeared not to be restricted
to certain months since there were some shifts
in peaks in separate years, suggesting a
response triggered by climatic factors.

25

100

50

AM J J A S ON D J F MAM J J A S ON D J FM
2008
2009
2010
Month

Fig. 3. Seasonal trends of monthly mean

reproductive litterfall. Vertical bar stands for the
standard error of the mean

-S Sharma et al.

26

ACKNOWLEDGMENTS
The preliminary version of the paper was
presented at the International Conference on
Environmental Aspects of Bangladesh (ICEAB
10) [Paper Code FM 01]. We are grateful to
our colleagues, Drs. S.M. Feroz, R. Suwa and
W. Min, and Messer. R. Deshar, for their
cooperation and active participation in the
field work. This study was financed in part by
a Grant-in-Aid of Scientific Research (No.
20510011) from the Ministry of Education,
Culture, Sports, Science and Technology,
Japan.

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2011, International Journal of Environment, 1(1): 2127