International Journal of Botany and

Research (IJBR)
ISSN 2277-4815
Vol. 3, Issue 1, Mar 2013, 19-28
© TJPRC Pvt. Ltd.

REPRODUCTIVE PHENOLOGY OF AEGICERAS CORNICULATUM (L) BLANCO- A

MANGROVE SPECIES IN THE GULF OF KACHCHH, GUJARAT, INDIA

PANDEY R1 & PANDEY C. N2

1
Project Coordinator-Climate Change Studies, GIPL, Gujarat Forest Department, Aranya Bhavan,
Gandhinagar, Gujarat, India
2
Principal Chief Conservator of Forest, (Wildlife), Aranya Bhavan, Gandhinagar, Gujarat, India

ABSTRACT

The reproductive phenology of A. corniculatum (L) Blanco has not been examined comprehensively in the Gulf of
Kachchh, Gujarat, India. The present work has examined the temporal and spatial variations in the reproductive phenology
of this species in this area. Besides, the study also reports the reproductive vigour (inflorescence production per tree) and
reproductive success of this species. The species remains in the reproductive phase throughout the year. The reproductive
cycle initiates during September- October and terminates during September of the next calendar year. The reproductive
phenology and reproductive vigour have shown temporal as well as spatial variations. The reproductive success has been
found to be different at different locations.

KEYWORDS: Mangrove, A. Corniculatum, Reproductive Phenology, Vigour

INTRODUCTION

The phenological studies have been carried out in various ecosystems throughout the world. These studies largely
tend to find relation in the phenological processes with biotic and abiotic factors (Frankie et al., 1974; Wielgolaski, 1974;
French & Sauer, 1974; Malaisse, 1974; Monasterio & Sarmiento, 1976; Tomlinson, 1986; Boinski & Fowler, 1989;
Seghieri et al., 1995; Morellato & Leitão-Filho, 1996; Brooke et al., 1996).

The phenology of mangrove forests has also been studied in various regions of the world (Gill & Tomlinson,
1971; Wium-Andersen and Christensen, 1978; Duke et al., 1984, Primack, 1985; Tomlinson, 1986; Duke, 1990; Marcus,
1999; Tyagi, 2003).

These studies largely pertain to the productivity of mangrove forests and its seasonal variations with reference to
the environmental conditions. Clarke (1995) has examined the growth and reproduction of A. corniculatum and A. marina
in the mangrove forests of New South Wales. Clarke, (1995) has also examined the population dynamics of this species in
the same mangrove forests.

The environmental conditions of a place are known to influence the phenological patterns (Liebermann, 1982).
Therefore, the phenological cycle tends to vary and, in turn, the reproductive cycle proceeds in accordance with the local or
regional environmental conditions. In spite of the relevance of the phenological events in the mangrove system, very few
authors have dealt with the phenological aspects of mangroves. As far as mangrove forests of Gulf of Kachchh, Gujarat,
India are concerned, no previous studies have been carried out about the reproductive phenology.

The present study aims at examining the temporal and spatial variation in reproductive phenology, reproductive
success and the spatial variations in the reproductive vigour of A. corniculatum in the Gulf of Kachchh, Gujarat.
20 Pandey R & Pandey C. N

MATERIALS AND METHODS

Study Area
The present work has been carried out at three islands viz. Pirotan -22°35'849'' North and 69°57'486" East-
(Population I), Sanada -22o34' 084'' North and 69o57' 374'' East -(Population II) and Patthapir -22o31'838'' North and
69o56'187'' East -(Population I) (Figure1.1) in the Gulf of Kachchh which falls under semi arid condition with average
annual rainfall of about 400 mm (Pandey, et. al., 2005). The average tidal amplitude is 5 m that may go even up to 8-10 m.
Most of the mangrove areas, therefore, get regular inundation.

Figure 1: Location of the Study Area in the Gulf of Kachchh, Gujarat

Reproductive Phenology- Crop Level

Ten matured trees were selected from each of the three research sites (30 trees from the three sites). These trees
were allotted specific Plant Index Numbers (PIN) and the range and the mean values of the heights and canopy perimeters
of these trees are indicated in Table 1.

Table 1: Range and Mean Height and Canopy Cover of Selected Trees

Height (m) Canopy Perimeter (m)

Site
Range Mean±SD Range Mean±SD
Patthapir 0.6 to 2.1 1.5±0.3 0.5 to 4.3 2.3±1.1
Pirotan 0.7 to 2.0 1.4±0.3 0.6 to 4.7 2.0±1.0
Sanada 0.8 to 2.0 1.3±0.9 0.2 to 2.8 0.9±0.5
SD: Standard Deviation

The inflorescences were categorized into four developmental stages viz. A-1, A-2, A-3 and A-4 (Table 2).
Reproductive Phenology of Aegiceras corniculatum (L) Blanco- A Mangrove Species in the Gulf of Kachchh, Gujarat, India 21

Table 2: Description of the Four Developmental Stage of A. Corniculatum

Duration of the
Developmental Characters Adopted for
Photograph Developmental Stage
Stage Identifying Different Stages
Represents

A-1 Inflorescences with buds only Maturation of buds

Inflorescences with flowers, bud

A-2 Flowering
and developing fruit (s)

Inflorescences with only

A-3 Maturation of fruit
developing fruits

Inflorescences with matured

A-4 Dispersal of fruits
propagules

The occurrence of these inflorescences was examined throughout the year for each population. For this, all the
inflorescences of the four developmental stages of selected trees were recorded fortnightly from April 2007 to May 2009.
Thus the observations pertains to 52 fortnights.

Reproductive Phenology-Inflorescence Level

For inflorescence level examination, 200 inflorescences of A-1 developmental stage were selected at each site
(600 inflorescences considering all the three sites) in October 2007. These inflorescences were observed fortnightly till the
surviving ones transformed into A-4 developmental stage. Hence, for each month two observations, corresponding to two
fortnights, were made. For instance, the two observations of January represented by Jan.1 and Jan.2 corresponding to the
periods of 1st to 15th January and 16th to 31st January respectively.

The A-4 developmental stage represents the inflorescence with all matured propagules only. During field
observations it was found that the flowering is asynchronous at inflorescence level. Therefore, within an inflorescence, the
individuals attain maturity at different time and as one individual attains the maturity, it gets dispersed. Since the
observations were made fortnightly, it was not possible to accurately note when all individuals of an inflorescence attained
maturity. Hence, the life of the inflorescence during A-4 developmental stage could not be estimated.

Reproductive Vigour

The inflorescence production per tree has been taken to represent the reproductive vigour of the three populations.
This information was deduced from the data collected to examine reproductive phenology at crop level.

Reproductive Success

This information was deduced from the data collected for inflorescence level examination of reproductive
phenology.

OBSERVATIONS
Reproductive Phenology-Crop Level
The reproductive phenology of A. corniculatum at the three locations i.e. occurrence of the four developmental
stages over the 52 fortnights are shown in Figure 2, 3, 4 respectively. The initiation of reproductive penology i.e. beginning
of the occurrence of A-1 developmental stages i.e. has been reported during September-October for all the three
populations. Further, the peak of A-1 stage was reported during December-January in all the three populations.
22 Pandey R & Pandey C. N

Figure 2: Reproductive Phenology of A. Corniculatum at Patthapir (Population I)

Figure 3: Reproductive Phenology of A. Corniculatum at Pirotan (Population II)

Figure 4: Reproductive Phenology of A. Corniculatum at Sanada (Population III)

The A-1 stage terminated during March in all the three populations during the reference years except at Pirotan
during 2009 when it terminated in April. Hence, the inflorescence with developing buds may be seen during September to
March in the Gulf of Kachchh. Further, the flowering which is represented by A-2 developmental stages have been
reported to initiate during October-November in all the three population during reference years except at Patthapir where it
Reproductive Phenology of Aegiceras corniculatum (L) Blanco- A Mangrove Species in the Gulf of Kachchh, Gujarat, India 23

initiated during December in the year 2008 (Figure 2). The flowering attained its peak during February-March in all the
populations. The flowering was reported to terminate during March- April in all the three populations. The developmental
stage A-3 (fruit development) representing the fruit development initiated December onwards and terminated during July
(except at Sanada in 2007). The peak of A-3 was attained -considering all the 52 fortnights of observation- during April-
May at Pirotan, April at Patthapir and May at Sanada. The developmental stage A-4 representing propagule maturation has
shown variation in its initiation for all the three populations. In the case of Patthapir (Population I), A-4 initiated during
Jun.1 (2007), May 2 (2008) and Apr.1 (2009). At Pirotan (Population II), the stage A-4 commenced in Jul.1 (2007), Jun.1
(2008) and Mar.1 (2009). At Pirotan, some A-4 developmental stages were also seen in Feb.1 during 2008. However at
Sanada, it initiated during Apr.2 (2007), May2 (2008) and Mar.2 (2009). The A-4 developmental stage was reported to
terminate by September in all the three populations, irrespective of their initiation periods (Figure 2, 3 and 4). Hence,
dispersal of matured propagules (A-4 developmental stage) complete by the end of monsoon season of the study area.

Reproductive Phenology - Inflorescence Level

The inflorescence life of the three developmental stages (A-1, A-2 and A-3) is summarized in Table 3. The
inflorescence life of the developmental stage A-1 and A-3 were similar and were significantly large that that at A-2 stage.
Among the three populations, the inflorescence life of all the three developmental stages was found to be longest at Pirotan
followed by Patthapir and Sanada.

Table 3: Inflorescence Life at Different Developmental Stages

Developmental Inflorescence Life (Mean ± s.e.) (Days)

Stage Pirotan Patthapir Sanada All Populations
A-1 117.9±3.82 110.6±1.60 106.3±2.10 110.4±1.31
A-2 19.6±1.18 15.4±0.73 14.0±0.51 15.6±0.43
A-3 117.4±2.94 106.8±1.70 104.6±1.85 108.9±1.34

Reproductive Vigour

The reproductive vigour has shown spatial and temporal variations. Among the three populations, the
reproductive vigour (inflorescence production per tree) was found to be the maximum at Pirotan followed by Patthapir and
Sanada (Figure 5).

Figure 5: Reproductive Vigour (Inflorescence Production Per Tree) of the Three Populations During 2007 to 2009
24 Pandey R & Pandey C. N

Further, it was reported to be higher during 2008 and relatively lower during 2007 and 2009. Hence, the
inflorescence production per tree has been found to show noticeable variation over time and space. However, the trends
were found to be similar.

Reproductive Success

Out of the 200 inflorescences, at each research site, 131 inflorescences at Pirotan island, 141 inflorescences at
Patthapir island and 78 inflorescences at Sanada island were found to have detached before attaining maturity of A-4 stage
as shown in Table 3.

Table 3: Reproductive Success of A. Corniculatum in three Populations

No. of Inflorescences
No. of No. of %
that Attained the
Population Selected Inflorescences Detached Prior Reproductive
Maturity of A-4
Inflorescence to Attaining Maturity of A-4 Success
(Reproductive Success)
Pirotan 200 131 69 34.5
Patthapir 200 141 59 29.5
Sanada 200 78 122 61

The reproductive success was found to be 34.5%, 29.5% and 61% in the populations of Pirotan, Patthapir and
Sanada. Considering all the three populations, the reproductive success comes out to be 41.66%.

DISCUSSIONS

The phenology of A. corniculatum has been studied by various scientists for different mangrove forests of the
world (Carey & Fraser, 1932; Duck et. al., 1984; Hutchings & Saenger, 1987) and for India (Singh, 1994; Kothari &
Singh, 1998; Naskar & Mandal, 1999; Banerjee et. al., 2002; Debnath, 2004). The flowering season of A. corniculatum
reported from various mangrove forests of India and that of Australia show noticeable variations as indicated in Table 4.
The flowering season of A. corniculatum has been reported to be different at east and west coasts of India. In fact,
flowering seasons were reported to vary even at single coast. In the case of Gujarat, Singh (1994) has reported the
flowering season of this species from January to February (2 months). However, the present work has reported the
flowering period from October to April (considering all the three populations together) with its peak during February-
March. It appears that Singh (1994) has reported the peak flowering period. It may be mentioned that the reproductive
phenology has shown temporal as well as spatial variations even in areas closely located (viz. the three populations of the
present study). Hence, the temporal and spatial variations in reproductive phenology is expected among locations separated
by larger distances.

Table 4: Flowering Season of A. Corniculatum Reported by Various Scientists

Researcher Study Area Flowering Season

Duck, 2006 Queensland, Australia January to September
Banerjee, 1989 India April to September
Banerjee ,2002 India February to April
East Coast of India
Debnath, 2004 India-Andaman & Nicobar March to September
Kumudranjan Naskar, 1999 West Bengal. Orissa January to September
West Coast of India
Kothari & Singh, 1998 Goa, Maharashtra, Gujarat February to October
Singh, 1994 Gujarat January to February
Present work Gujarat October to April
Reproductive Phenology of Aegiceras corniculatum (L) Blanco- A Mangrove Species in the Gulf of Kachchh, Gujarat, India 25

Clarke (1994, 1995) has given detailed information about the reproductive phenology of Aegiceras corniculatum
of Queensland forests, Australia. The reproductive phenology of A. corniculatum reported by Clarke (1994) for Australian
mangrove forests and that reported for mangroves of the Gulf of Kachchh, Gujarat, India during the present study are
summarized in Table 5. Clarke (1994) reported that the commencement of the bud formation occurs during May and
continues up to November. In the present study, this (stage A-1) was reported during September to April with peak in
December-January.

The flowering was reported during July to November in the Queensland mangroves (Clarke, 1994). However, in
the case of Sydney mangrove forests (Carey & Fraser, 1932) earlier flowering was reported. In the present work, flowering
has been reported from October (beginning to winter) to April (beginning of summer) with peak during February-March.
Further, Clarke (1994) reported commencement of flowering during the winters (July) with a peak during the summer
(December). Clarke (1994) has reported immature fruits (A-3) during December to March. In the case of the present work,
it was found to be during December to July. Further, the present work has reported matured fruits (A-4) during March to
September and Clarke (1994) reported the same during March-April for the Australian mangrove forests.

Table 5: Reproductive Phenology of A. Corniculatum in Indian and Australian Mangrove Forests

Australia (Clark 1994) India, Gujarat (Present Work)

Reproductive *Duration of **Life Span of that
Duration
Phenology Period Period Occurrence Developmental
(Months)
(Months) Stage (Months)
Initiation and
May to September to
continuation of bud 7 8 About 4.0
November April
formation (A-1)
July to October to
Flowering (A-2) 5 7 About 0.5
November April
Fruit formation December to December to
4 5 About 4.0
(A-3) March July
March to
Fruit maturation (A-4) March-April 2 7 About 0.5
September
Total Duration of
About 12-14 Months About 12 Months About 9 Months
Reproductive Cycle
* Occurrence of a Particular Developmental Stage Over the Period of 12 Months and ** Inflorescence Life of a
Particular Developmental Stage

It has been reported that the reproductive phenology of A. corniculatum is influenced by the distance from the
tropical latitude (Duke, 1990). Duck (2006) has reported that the period between flowering and fruiting increased from 2-3
months in northern tropical areas to 10 months in southern temperate areas. This is because the growth rate of reproductive
development increases two to three times with the increase in temperature of 100C each (Duke, 1990). Though Clarke
(1994) has not mentioned whether the duration of various phenophase (developmental stages) represents the inflorescence
level occurrence or crop level occurrence, it appears that it represents crop level occurrence (probably peak occurrences at
crop level).

The present study has also reported that the period between peaks of flowering and fruiting is 2 to 3 months in the
study area which falls close to the Tropic of Capricorn. Thus, the observations of the current study are in agreement with
Duck (2006) for areas close to tropics. Further, the overall duration of the reproductive phenology is similar in the
mangrove forests of Queensland (12 to 14 months) and Gulf of Kachchh (12 months). However, the occurrence of each
developmental stage is spread over larger period in the Gulf of Kachchh, Gujarat. Though the inflorescence life is only
about 9 months, the reproductive phenology is spread over 12 months in Gulf of Kachchh. The reproductive vigour of a
26 Pandey R & Pandey C. N

different mangrove species-Rhizophora stylosa, Rhizophora mangle (synonym R. samoensis) and Bruguiera gymnorhiza
has been reported to be higher in wet areas than those of dry areas (Tyagi, 2003). However, the reproductive vigour of A.
corniculatum has not been examined. The reproductive vigour of the three populations of A. corniculatum have shown
noticiable varations during the current study. In the case of A. corniculatum, a significant number of inflorescences get
detached prior to attaining maturity as well developed propagules. Considering all the three populations, only about
41.66% of the total inflorescences were found to achieve reproductive success and the remaining 58.34% samples were
observed to have detached prior to achieving maturity of fruiting (A-4). High pre-dispersal mortality (up to 95%) in A.
corniculatum has been reported from other mangrove areas of the world (Clarke, 1995).

ACKNOWLEDGEMENTS

We acknowledge GEER Foundation, Gandhinagar for providing necessary support for implementing the project.
We also acknowledge the Ministry of Environment and Forest (MoEF), Gov. of India for sponsoring the project and to
Gujarat Forest department for providing the local help and the permissions for field works. The thanks are also due to Dr.
Harshad Salvi (Research associate, GEER Foundation), Mrs. Urvi Bhatt (Senior Research Fellow, GEER Foundation), Ms.
Anjali Sharma (Senior Research Fellow, GEER Foundation), Mr. Shailesh Dodiya (Senior Research Fellow, GEER
Foundation), Mr. Nilesh Vaghela (Senior Research Fellow, GEER Foundation), Mr. Sandeep Patel (Junior Research
Fellow, GEER Foundation), Mr. Irshad Theba (Research assistant, GEER Foundation) for helping field observations. We
are also thankful to the boatman Mr. Junusbhai and his team.

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