Professional Documents
Culture Documents
135-144, 2010
2010 WIOMSA
Kenya Marine and Fisheries Research Institute (KMFRI), P. O. Box 81651, Mombasa,
2
Kenya; 3Biocomplexity Research Focus c/o Laboratory of General Botany and Nature
Management, Mangrove Management Group, Faculty of Sciences, Vrije Universiteit Brussel
- VUB, Pleinlaan 2, B-1050 Brussel, Belgium; 4Biocomplexity Research Focus (Complexit
et Dynamique des Systmes Tropicaux), Dpartement de Biologie des Organismes, Facult
des Sciences, Universit Libre de Bruxelles - ULB, Campus du Solbosch, CP 169, Avenue
Franklin D. Roosevelt 50, B-1050 Bruxelles, Belgium.
300 34
250 32
Temperature(oC)
Rainfall(mm)
30
200
28
150
26
100 24
50 22
0 20
J F M A M J J A S O N D
Months
Fig. 2: Monthly rainfall (mm) and mean monthly temperatures (C) at Mombasa in 2005-2006. April-
June comprises the long rain season and October-November the short rain season. (Source: Mombasa
Meteorological Station.)
138 V. W. WANGONDU ET AL
Table 2: Mean, minimum and maximum longevity (months) of A. marina leaves at sites in Gazi Bay.
Ns = natural site; rs = reforested site; n = number leaves that emerged and fell during the study period.
Site description n Mean(S.E) Minimum Maximum
1 (Landward ns) 278 11.22 (4.73) 0.93 18.66
2 (Landward ns) 292 10.56 (3.88) 1.40 16.80
3 (Landward ns) 320 10.95 (3.67) 0.46 17.73
4 (Seaward ns) 124 10.63 (4.07) 1.86 18.66
5 (Seaward rs) 233 9.24 (4.04) 0.93 16.80
Trends in reproductive phenology fruit production between the sites. Peak fruit
production occurred in April and May which
The timing of the reproduction cycle in
coincided with the wet season. It is notable
A. marina was seasonal and its initiation
that reproductive activity in 2005 was lower
coincided with the short rainy season. Bud
at all sites compared to 2006 and, though bud
initiation started in the month of November
production was recorded at all the sites in 2005,
at all A. marina sites during the study period.
fruits were only found at the reforested site.
Peaks in bud production were observed in
December and January at all the sites (Fig. 3 Leaf longevity
and 4). However, no new buds were produced
after February at the landward sites, whereas Mean leaf longevity in A. marina was 11
bud formation was prolonged at the seaward months (Table 2). There was a significant
sites until April and March in 2005 and 2006 difference in leaf longevity between the sites
respectively (Fig.3). Bud production was (p<0.001), with the seaward reforested site
much lower at all sites in 2005 than in 2006. manifesting the shortest leaf longevity of
Bud production also differed significantly 9.24 4.04 months and site 1 the greatest leaf
among sites (p<0.001). Landward site 1 and longevity of 11.22 4.73 months
seaward site 4 differed significantly from
landward sites 2 and 3 (p<0.01) as they DISCUSSION
produced the least number of buds.
At the landward sites, flower production Vegetative phenology
commenced in November to March in site Seasonality was observed in A. marina leaf
2 and December to March in sites 1 and 3 production and fall at both the landward and
(Fig. 3). It started later at the seaward sites, seaward sites but the period of leaf production
from January to March in site 4, and January was longer at the former (January-April)
to May in site 5 (Fig. 4). Flower production in 2005. Thus the vegetative phenology of
was higher in 2006 than in 2005 but lower landward and seaward A. marina varied with
than bud production at all the sites. There respect to the timing and abundance of leaf
was a significant difference (p<0.001) in emergence and fall.
flower production between sites, with site 4 Unimodal and bimodal peaks in mangrove
producing the least flowers and having the leaf production are known to be associated with
shortest flowering period of three months. forests growing in higher and lower intertidal
Avicennia marina fruits were observed areas respectively due to fluctuations in ground
from the months of March to May at the water salinity (Wium-Andersen, 1981). High
landward sites with a peak in April. At the interstitial water salinity elevates stress in
seaward sites, fruits were observed from mangroves, resulting in increased leaf loss to
March to May in site 5 and July in site 4. There reduce water loss from transpiration, especially
were no significant differences (p>0.05) in during the dry season, thus reducing water
140 V. W. WANGONDU ET AL
(Site 1) Leaf fall Leaf em ergence Buds Flow ers Fruits
(Site 1) Leaf fall Leaf em ergence Buds Flow ers Fruits
(Site6 1) Leaf fall Leaf em ergence Buds Flow ers Fruits
0.35
(Site 1) Leaf
(Site 1) 6 Leaf fall
fall Leaf
Leaf em
em ergence
ergence Buds
Buds Flow
Flow ers
ers Fruits 0.35
Fruits/Shoot
6 0.30
0.35
Buds/Shoot
65 0.35
0.25
Mean Leaf fall &
Fruits/Shoot
4 0.30
Fruits/Shoot
Buds/Shoot
5 0.25
Fruits/Shoot
& &
0.30
Buds/Shoot
5
54 0.30
Buds/Shoot
0.20
&&fall
0.25
3 4 0.20
0.25
fall
0.25
Leaf
4
43
fall
0.15
0.20
fall
Flowers,
0.15
0.20
emergence,
Leaf
2 3 0.20
Leaf
Flowers,
Mean
3
Leaf
32 0.15
0.10
emergence,
Flowers,
0.10
Flowers,
0.15
emergence,
Mean
0.15
emergence,
2
Mean
1 2 0.10
Mean
21 0.05
Mean
0.05
0.10
0.10
1
Mean
1 0.05
0 0.00
Mean
10 0.00
0.05
Mean
0.05
J 0 F JMF AM MA J M JJ A J AS SO ON NDDJ J FF MMAA M
M J JJ AA SS OO NND D0.00
0
0 J F M A M J J A S O N D J F M A M J J A S O N D 0.00 0.00
J
J F F MM A M
M J
A 2005 J
J A
2005
J A SS OO N
N DD J
J F F MM A
A M
M J J
J A
2006
2006
J A SS O
O N N DD
2005 2006
2005
2005 2006
2006
(Site 2)
(Site 2)
(Site 2)
(Site
(Site 2)
2)
10 1.2
10 10 1.2
Fruits/Shoot
1.2
10 1.2
emergence,Buds/Shoot
1.0
Fruits/Shoot
emergence,Buds/Shoot
1.01.0
Fruits/Shoot
emergence,Buds/Shoot
& &
Fruits/Shoot
8
emergence,Buds/Shoot
1.0
0.8
emergence,Buds/Shoot
&&fall
8 1.0
Mean Leaf fall &
86
0.80.8
fall
Leaf
6 0.8
0.6
fall
6 0.8
fall
Flowers,
6
Leaf
64 0.60.6
Leaf
Mean
Flowers,
0.6
Leaf
0.4
0.6
Flowers,
4 44
Flowers,
Mean
42 0.40.4
Mean
0.4
Mean
0.2
0.4
Mean
2 0.2
2 2
Mean
20 0.00.2
0.2
Mean
0.2
0 J F M A M J J A S O N D J F M A M J J A S O N D 0.0 Mean
0 0 0 0.00.0
J F M A M J J A S O N D J F M A M J J A S O N D 0.0
J FJ J MF
FAMM
M AM
A M J J
2005
J JJ J
AAAS S
SOO
ONNNDD
DJJJ FF
FMMA
M A M
M
MJ J JJ A
2006
J J AA S
SSO
OONN ND
DD
2005 2006
2005
2005 2006
2006
(site 3) 2005 2006
(site 3)
(site 3)
(site 3)
(site 3)
16 2.0
Flowers,Fruits/Shoot
16 2.0
emergence,Buds/Shoot
16
16 2.0
2.0
1.6
Flowers,Fruits/Shoot
16 12 2.0
Flowers,Fruits/Shoot
emergence,Buds/Shoot
& &
Flowers,Fruits/Shoot
1.6
emergence,Buds/Shoot
emergence,Buds/Shoot
&&fall
Mean Flowers,Fruits/Shoot
12 1.6
1.6
12 1.2
emergence,Buds/Shoot
12
fall
1.6
Leaf
fall
12 8 1.2
fall
Mean Leaf fall &
1.2
Leaf
8 1.2
0.8
Leaf
Mean
8
Leaf
8 0.81.2
4
Mean
0.8
0.8
8 0.4
Mean
Mean
Mean
4
4
4 0.40.8
Mean
0.4
0.4
0 0.0
Mean
Mean
4 J
J J A S O N D J F M A M
0 F M A M J J A S O N D 0.00.4
0
0 0.0
J
J J A S O N D J F M A M F M A M J J A S O N D 0.0
J J
J A
J2005 A S
S OO NN D D J J
J F
J F M F
M A
FA MM
M
M A
A M
M J J
J A
J2006 A S S OON D
N D
Fig 3:0 Leaf fall and emergence,
2005
production
2005 of buds, flowers and 2006 in A. marina at the 0.0
fruits
2006 landward
Sites 1, 2 JandF3 M J 2005
A MBay.
in Gazi J A S O N D J F M A M 2006
J J A S O N D
2005 2006
Phenology of Avicennia marina (Forsk.) Vierh. in a Disjunctly-zoned Mangrove Stand in Kenya 141
(site
(site4)
4)
Leaf
Leaffall
fall Leaf
Leafemergence
emergence Buds
Buds Flowers
Flowers Fruits
Fruits
(site 4)
Leaf fall Leaf emergence Buds Flowers Fruits
Fruits/Shoot
&&Fruits/Shoot
66 0.08
0.08
emergence,
fall&&emergence,
55
0.06
0.06
& Fruits/Shoot
6 0.08
Buds/Shoot
44
Buds/Shoot
& emergence,
35
3 0.04
0.04
Flowers
0.06
MeanFlowers
fall
24
Buds/Shoot
2
Leaf
0.02
fallLeaf
0.02
13
1 0.04
Mean Flowers
Mean
Mean
Mean
2
00 0.00
0.00
0.02
Mean Leaf
1 JJ FF M
M AA M
M JJ JJ AA SS OO NN DD JJ FF M
M AA M
M JJ JJ AA SS OO NN DD
0 2005
2005 2006
2006 0.00
J F M A M J J A S O N D J F M A M J J A S O N D
2005 2006
(site
(site5)
5)
66 0.5
0.5
Fruits/Shoot
&&Fruits/Shoot
(site 5)
emergence,Buds/Shoot
55
emergence,Buds/Shoot
0.4
0.4
6 0.5
fall&&
& Fruits/Shoot
44
& fall
0.3
emergence,Buds/Shoot
5 0.3
0.4
Leaf
fallLeaf
33
Flowers
MeanFlowers
4 0.2
0.2
Mean
Mean
22 0.3
Mean Leaf
3
0.1
Mean Flowers
11 0.1
0.2
Mean
2
00 0.0
0.0
1 0.1
JJ FF M
M AA M
M JJ JJ AA SS OO NN DD JJ FF M
M AA M
M JJ JJ AA SS OO NN DD
2005
2005 0 2006
2006 0.0
J F M A M J J A S O N D J F M A M J J A S O N D
Fig 4: Leaf fall and emergence, production of buds, flowers and fruits in A. marina at the seaward
Sites 4 and 5 in Gazi Bay. 2005 2006
stress. Increased leaf loss by the landward A. little vegetative activity while the trees were
marina trees during the dry season is indicative flowering and fruiting, and leaf production
of their adaptive measures to reduce water loss. and fall only resumed in June after completion
The lack of clear unimodal and bimodal leaf fall of fruit fall. In 2005 there was little or no
and emergence patterns at both the landward reproductive activity and it was associated
and seaward sites in 2006 compared to 2005 with higher leaf fall and emergence. These
may be attributable to the high and continuous results are corroborated by other studies
rainfall experienced in that year. This may have (Duke et al., 1984; Wium-Andersen and
lowered the salinity levels of the interstitial Christensen, 1978; Coupland et al., 2005).
water, thus reducing salt stress and changing Increased leaf production by A. marina
the leaf fall and emergence patterns. has been reported during the wet season in
The lower leaf fall and emergence in 2006 Darwin Harbour in Australia (Coupland et
may also be related to the reproductive status al., 2005). Ochieng and Erftemeijer (2002)
of the trees at that time. There possibly was reported unimodal leaf flush patterns in
a link between vegetative production and landward A. marina at Gazi in June to July (wet
the reproductive cycle in that there was very season) in 1993-1994. In the present study,
142 V. W. WANGONDU ET AL
similar observations were made between dry months which corroborates the results of
July and September, which is normally a dry other studies (Duke et al., 1984: Fernandes
season, indicating the likelihood of a shift in 1999; Ochieng & Erftemeijer, 2002).
phenological trends in species with respect to Avicennia marina in Gazi Bay (4 25S, 39
the prevailing climatic conditions. Peak leaf 3 E) was nevertheless observed to flower
fall occurred one to two months after peak later than in Darwin Harbour (12 26S, 130
leaf emergence at all the sites, corresponding 51 E) (Coupland et al., 2005).
with findings on A. marina in Australia (Duke, Significant differences in bud and flower
1988). Shifting of leaf fall and emergence production between the sites can be related
peaks to later months, especially in 2006, to stand structure and tree height. Avicennia
indicates that the phenological patterns are marina trees at sites 2 and 3 were short and
not static and may be influenced by changing stunted and had a lower canopy cover, thus
climatic factors; this is evidenced by the allowing more light penetration due to less
different rainfall and air temperatures during shading. Sites 1, 4 and 5 conversely had taller
the two years of study. However, long-term trees and higher canopy cover. This may
phenological studies are necessary to draw explain why a greater production of buds and
conclusions regarding the effects of climate flowers was recorded at sites 2 and 3.
change on mangrove phenology. Timing of fruit production in the two zones
The leaf longevity in A. marina observed was uniform; however, fruits at the seaward
in this study corresponded with that reported site 4 persisted up to June. Fruit fall in the two
for the same species at Gazi Bay (Ochieng & zones coincided with the wet season, consistent
Erftemeijer, 2002). It is close to the 14 months with findings in other studies (Duke et al.,
reported for the same species in Australia 1984; Fernandes, 1999; Ochieng & Erftemeijer,
(Coupland et al., 2005). Significant differences 2002; Coupland et al., 2005). The reproductive
in leaf longevities at the landward and seaward cycle of A. marina from bud initiation to fruit
sites may be indicative of a response to fall was observed to last 6-7 months in the
stress brought about by differences in ground landward zone and 7-8 months in the seaward
water salinity resulting from differences in zone. Other studies have reported varying
inundation. The seaward reforested and natural reproductive cycles at different latitudes, e.g.
sites in this study manifested reduced leaf five months in Darwin Harbour (Coupland et
lifespan. Frequent inundation of the seaward al., 2005), six months in southern Thailand
sites results in less stress, hence possibly a (Wium-Andersen & Christensen, 1978) and
higher leaf turnover. The seaward reforested one year in south-eastern Australia (Clarke &
site 4 comprised a young stand (12 yrs old) Myerscough, 1991). Zonation may have had an
that may have been actively growing, causing influence in fruiting of A. marina since sites that
a similar reduction in leaf longevity. were frequently inundated (seaward) supported
prolonged budding and fruiting relative to
Reproductive phenology those that were less inundated (landward sites).
The synchronised initiation of budding at all The relationship between vegetative and
sites may be indicative of the adaptability A. reproductive cycles cannot be ignored in this
marina in the two zones. However, earlier study as reproductive months were associated
flowering of trees in the landward zone than with very little vegetative activity, both in terms
those in the seaward zones was indicative of of leaf production and fall. Leaf production
a lack of uniformity in phenological trends peaked prior to bud initiation and again later
in this species, even within the same locality. after fruit fall in May. This corroborates the
Prolonged flowering at the seaward sites work of Duke (1990) who found a link in the
may have resulted from more favourable timing of leaf appearance and leaf fall with
environmental factors compared to the inflorescence development. This has been
landward sites. Flowering peaked during the attributed to resource partitioning within plants
Phenology of Avicennia marina (Forsk.) Vierh. in a Disjunctly-zoned Mangrove Stand in Kenya 143