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Hikobia 15: xx–xx.


Does mangrove Kandelia candel (L.) Druce follows a mangrove zona-

tion, soil salinity and substrate for survival?

ROTAQUIO, JR., E. L., NAKAGOSHI, N. & ROTAQUIO, R. L. 2008. Does mangrove Kan-
delia candel (L.) Druce follows a mangrove zonation, soil salinity and substrate for
survival? Hikobia 15: xx–xxx.

The question why mangrove plants are forming mangrove zonation or distinct divi-
sion or separation among each species in the ntertidal flat has been the subject of
arguments among mangrove scientists and researchers. How zonation affects the
survival of a certain mangrove is still unclear. It is in this light that this study was
conducted to verify the significance of mangrove zonation to the species of Kandelia
candel (L.) Druce. The findings showed that this species has a higher chance of sur-
vival if planted in its preferred zone with due consideration to its required soil salin-
ity and soil substrate type. This study also revealed that mangrove zonation is a natu-
ral pattern to follow or as a basis of what type of species should be planted in order to
attain a higher planting survival in the mangrove reforestation activity.

Eutiquio L. Rotaquio, JR., Graduate School for International Development and Coop-
eration, Hiroshima University, Kagamiyama, Higashi-Hiroshima, 739–8529, Japan;
Aurora State College of Technology Baler 3200 Aurora Province, Philippines. E-mail
Nobukazu Nakagoshi, Graduate School for International Development and Coopera-
tion, Hiroshima University, Kagamiyama, Higashi-Hiroshima, 739–8529, Japan. E-
mail Address:
Ronaldo L. Rotaquio, Provincial Environment and Natural Resources Office, Depart-
ment of Environment and Natural Resources, Baler 3200, Philippines. E-mail Address:

ward to landward.
1. Introduction
 There are several definitions of mangrove zo-
 One of the distinguishing characteristics of nations given by mangrove scientists and ecolo-
major mangrove plants is the formation of zona- gists. Primavera (2000) defined it as the presence
tion, of which each species from seaward zone of stands of one or two species depending on in-
going to the landward most part has its own ter-tidal position and location within an estuary.
distinct division of species association vary- According to Tomlinson (1986) the existence of
ing in different length and width parallel to the zone is often mono-specific and an impression
shoreline or river edge. The extent in terms of that it is a series of vegetational bands parallel to
wideness and length depend on the availability the coastline. Zonation could be defined as the
of the mudflat and the topography of the area. It structural feature of mangrove forest in some
can also be described as the distinctive formation parts of the world (Woodroffe 1992). The simple
of pure community of each species with visible explanation for mangrove zonation according to
boundary and growing parallel to the coastal wa- Hogarth (1999) is that, it represents a steady state
terline from lowest tide line to the maximum tide response to a gradient in some critical physical
line. It is the distinct separation of each major variable, and in the case of upshore/downshore
mangrove species in the inter-tidal flat from sea- gradients that this gradient is maintained by the
2 Hikobia Vol. 15, No. 2, 2008

tidal inundation regime. Zonation is modified uting factors in the formation of zonation may
by local topography, which determines tidal and include plant succession, geomorphology, physi-
fresh-water runoff, and by sediment composition ological adaptation, propagule size, seed preda-
and stability (Semeniuk 1980). tion, and interspecific interactions. However,
 The formation of mangrove species zonation these factors depend also on the individual habi-
has been the subject of several studies and argu- tat type or requirements of each species. On the
ments among ecologists arguing that the tropical other hand, Robertson and Alongi (1991) argued
mangrove forest is characterized by clear zona- that plant succession plays a minor role in man-
tion along tidal gradient, and it has been sup- grove zonation and that simple erosion and sedi-
posed that the zonation is primarily controlled mentation control the distribution of mangroves
by soil factors. Hogarth (1999) stated that there along the seaward edge of the mangal.
are four main approaches to the causes of spe-  Tidal regime or current also affects the forma-
cies zonation which have emerged based on the tion of some mangrove zonations such as in In-
concepts of population dynamics, ecophysiology, dian Sunderbans, the mangal areas that regularly
and geomorphology. The first one is that, zona- inundates by tide diurnally is dominated by Avi-
tion results from the physical sorting of floating cennia marina and Avicennia alba while Excoe-
propagules by water movement, the position of caria agallocha, Ceriops decandra and Acanthus
an adult tree within a forest being determined by ilicifolius dominate sites that are not completely
the point at which a propagule was stranded. Sec- indundated (Saha & Choudhury 1995).
ond, selection might take place after settlement,  Tomlinson (1986) stated that there are actually
(e.g. different species thriving at different posi- 35 interconnecting environmental variables in
tions along some physical gradient). Third, zona- determining vegetation zonation. However, these
tion might be a consequence of gradients created variables might be contrasted on the basis of
by major geomorphological changes and finally, whether the influence is abiotic or biotic. Major
zonation might be the product of ecological in- abiotic include geomorphology, tidal inundation
teractions between species in the community, classes and physiological responses to gradients
the sequence of species along a transect in space such as salinity. Biotic factors influencing zona-
corresponding to a natural succession of species tion include propagule sorting and competition.
with time. Rabinowitz (1978) in her observation Semeniuk’s (1980) interpretation of physiography
in Panama that the shore level at which a species and hydrology provides evidence that ground-
occurred correlated with propagule size, that the water salinities are an important influence on
species thriving in the landward most part of the mangrove distribution, that if there is great
mangrove were those with smaller propagules. ground-water seepage, there was a formation of
Larger propagules were more likely settled and landward fringe and the lack of ground-water
established at the seaward, which is regularly seepage, almost no formation of landward fringe
flooded. Kathiresan and Bingham (2001) stated of mangroves.
that one potential cause of mangrove zonation is  Based on the foregoing premises, it is impor-
the differential ability of propagules to establish tant to point out about what is the limiting fac-
at different tidal heights. According to them it is tor in the formation of mangrove zonation? The
directly related to propagule size. It has been sug- above-mentioned arguments are too general with
gested that small propagules drift further inland a wide range of scope and therefore requires ex-
and establish better in shallow water than large perimental works to answer. This study aimed
propagules however, this case is not true in the to answer some of the foregoing questions deal-
case of Avicennia marina, of which the seed is ing and limited only to the two most important
generally small but most likely appeared in sea- ecological factors affecting the formation of
ward mangrove where strong water current and mangrove zonation- the soil salinity and type
tide is relatively strong and high as in the case of of soil substrate. This particular study was also
mangrove in Aurora Province, Philippines where intentionally conducted to answer why most of
this study was conducted. the local and national mangrove reforestations
 Woodroffe (1992) pointed out that the contrib- in the Philippines were always failure. In the lo-

cal place of Aurora, Philippines alone, five major (15°44′936″ N; 121°34′633″ E), Aurora, Philip-
reforestation projects have been implemented, pines. Thirty propagules were planted in Oc-
however, the average percentage survival was tober, 2005 in an open space in each zone in
45% and below (AIADP II 1988) far behind to the same mangrove forest, with spacing of 0.5
the national standard of 80% reforestation sur- m × 0.5 m. No other silvicultural treatment and
vival rate to be considered as successful one. management have been applied in the planted
Possible common mistake based on the actual propagules and assumed that the growth and
observation of the authors were the mismatched survival were just affected by natural factors ex-
of the kind of planted propagules in different isting in the mangroves. The planted propagules
zonations since most of the propagule used was were monitored and checked its survival month-
only one species, Rhizophora stylosa, for all ly, including the in situ soil salinity in each zona-
the zonations in the mangrove areas because of tions. The monitoring period lasted for one year.
abundant and available propagules. Aside from
this, common people who were involved in the 2.2 Methods of Monitoring the Survival of Kan-
reforestation strongly believed that any kind of delia candel
species of mangrove can survive everywhere in  Initially, prior to planting, each propagules were
mangrove forest because it is considered as halo- checked.  Damaged and deformed propagules
phyte plants, which means according to them, were eliminated. Only sound and vigorous
it has the ability to cope up or has the tolerance propagules with closed terminal shoot and/or leaf
with saline environment. However, in the opinion bud was chosen for the experiment. The basis for
of the author, this idea is too general and may survival was the continuous presence of terminal
provide half-baked information for mangrove bud and production of leaves until the end or ter-
researchers. To test this notion and to provide mination of observation period.
evidence, we conducted an experimental test for
the survival and adaptability of Kandelia can- 2.3 Determination of soil salinity
del (L.) Druce in different mangrove zonations.  In measuring soil salinity, the Twin Cond
This species is the latest major mangrove species Horiba Conductivity Meter B–173, which was
added to the list of existing mangrove species in calibrated with 1.41 standard solution into the
the Philippines and discovered existing naturally sensor cell following the 1:1 soil to water method
only in the mangroves of Aurora throughout the (Whitney 1998).  Three replicates of 20 g of soil
entire Philippine archipelago.  So far, no other samples were taken randomly from each zonation
records of its natural population in other suit- at 20 cm soil depth, and then, 20 mL of distilled
able mangrove areas in the Philippines have been water was added and mixed to the samples and
reported.  Our hypothesis is that each species allowed it to settle for 15 minutes.  The conduc-
of mangrove follows a zonation appropriate for tivity meter was then dripped into the suspension
them to survived, disregarding their required to get the conductivity readings.  The conductiv-
zonation, the type of substrate and its level of ity readings were then converted into salinity in
soil salinity requirements in planting, efforts in parts per thousand (ppt) unit of measurement.
reforestation activities might possibly fail.  This The monthly in situ soil salinity readings in dif-
study aimed to test the survival and adaptability ferent zones where Kandelia candel has been
of Kandelia candel in the different mangrove planted is presented in Table 1.
zonations with due respect to the soil salinity and
type of soil substrate. 2.4 Determination of mangrove zonation and
identification of mangroves
 To measure the extent of mangrove zonation,
2. Materials and methods
three transect lines in the whole mangrove for-
2.1 Plant materials and experimental site est were laid out perpendicular to the shoreline
 A total of 330 fallen and mature propagules or river edge ranging from 1–782 m using a steel
of Kandelia candel was collected around the transect tape from the lowest tide level line up to
vigorous mother trees in the mangroves of Baler the maximum tide level line.  The two transect
Table 1. In situ soil salinity readings (parts per thousand) in different zones where Kandelia cadel has been planted.

Table 2. Extent of the each zonations from seaward to landward in the mangrove areas of Baler, Aurora Philippines.
Hikobia Vol. 15, No. 2, 2008

lines were set at the two end parts of the man- with 4–55 m, Sonneratia alba zone with 5–25 m,
grove forest and the other one was set on the Rhizophora apiculata with 0–20 m, Bruguiera
middle of the forest all starting from seaward (mixed varieties) having a zone width of 0–200
going to the landward. It was followed by mea- m, Kandelia candel zone on the other hand can
suring the extent width of each zonation across be found along the river bank of Zabali and
the transect lines and then the mean average was sparsely distributed up to the small creek closed
computed for each zone. to the Aurora State College of Technology (AS-
 Species were identified using the dichotomous COT) Marine Laboratory.  The zonation width
key in mangrove identification (Tomlinson 1986), is relatively small ranging from 0–5 m and/or an
using portable monographs and other mangrove average of 1.6 m and growing mostly as frontal
reference books.  One dendrologist and one for- pioneer species in muddy river edge. Huge trees
ester joined in the fieldwork to make sure with of old growth Lumnitzera littorea having a zone
the identification and field data gathering. Sample width of 0–250 m can be found on the inner side
specimen for unidentified and species with doubt of mangrove forest.  This was followed by Heri-
identifications were sent to the University of the tiera littorea zone, with an average zone width of
Philippines experts for further verification and 5 m, Acanthus ebracteatus zone with 4–10 m and
proper identification of the species. Acrostichum aureum zone with small zone width
of 0–2 m located in the landward most and well
disturbed areas.  Extent of each zonation from
3. Results
seaward to landward in the mangroves of Baler,
3.1 Baler mangroves and its zonation Aurora Province, Phillippines is presented in
 The mangal areas of Baler has a total area of Table 2.
19.7 ha (AIADP II 1988).  The mangrove zona-
tions are represented by Rhizophora stylosa (arti- 3.2 Survival and adaptability of Kandelia candel
ficially planted) occupying the most seaward part in different mangrove zonations
with sandy to coralline soil substrate on the east-  There were 11 distinct zonations identified in
ern side of Baler port. Zone width ranges from Baler mangroves from seaward to landward most
0–250 m, followed by Avicennia marina zone part.  These include Rhizophora stylosa zone,

Fig. 1. Vertical profile of mangrove zonations at Baler, Aurora Province, Philippines.

6 Hikobia Vol. 15, No. 2, 2008

Table 3. Adaptability survival of planted Kandelia candel in different mangrove zonations.

Fig. 2. Percentage survival of Kandelia candel planted in different mangrove zonations at Baler, Aurora,

Avicennia marina zone, Sonneratia alba zone, test, there was no survival in two seaward most
Rhizophora apiculata zone, Bruguiera zone , zones (Rhizophora stylosa & Avicennia marina
Kandelia candel zone, Lumnitzera littorea zone, zones). In Rhizophora stylosa zone, all seedlings
Heritiera littoralis zone, Acanthus ebracteatus planted were already died after 7th months of
zone, Nypa fruticans zone and Acrostichum au- planting, worst case in Avicennia marina zone
reum zone.  Although the Heritiera littoralis, wherein no more surviving propagules just only
Acanthus ebracteatus and Acrostichum aureum after four months of planting. On the other hand,
are not considered as major mangrove based on four zonations were noted having highest per-
the criteria set by Tomlinson (1986), they formed centage survival.  These are the Bruguiera gymn-
a distinctive zonation in Baler mangrove and orrhiza zone with 57%, Nypa fruticans zone with
therefore we tried to include these zones in do- 63%, Rhizophora apiculata zone with 67% and
ing the experiment.  The vertical profile of Baler the highest was recorded from its original zone
mangrove zonations from seaward going to the (Kandelia candel zone) with 70%.  The adapt-
landward is presented in Fig. 1. ability percentage survival of planted Kandelia
 Based on the data gathered at the end of the candel in each zone for one year monitoring is
Table 4. Survey Details and Mangrove Species Present in Baler, Aurora, Philippines.
8 Hikobia Vol. 15, No. 2, 2008

numerically presented in Table 3 and graphically both riverine mangrove with muddy substrate
presented in Fig. 2. and coastal type of mangrove with coralline and
rocky substrate directly facing the Pacific Ocean
3.3 Soil substrate and usually exposed to strong tidal current and
 The type of soil for each zonation was gener- waves especially during bad weather conditions.
ally categorized into rocky or coralline, sandy, Although disturbed by such conditions, the spe-
muddy and muck based on the description given cies of Rhizphora stylosa is still surviving and
by Melana et al. (2000).  Rocky or coralline had formed a distinct zonation. The stilt or aerial
substrate is characterized as hard shelves where roots of this species lying in between the holes of
small or thin pockets of softer sediment and dead dead corals and rocks which make the stem more
coral reefs are found.  Sandy substrate comprises stable to withstand against strong tidal current.
of tiny grains of sediments e.g. coral usually less Following this zonation is the Avicennia marina
than 2 mm in diameter. There is no organic mat- zone, then, Sonneratia alba zone, which some-
ter to speak of, although it may overlay mud or times overlapped and mixed together with almost
muck. Like mud, sand may be as shallow as 2–3 the same substrate types combination of coral-
cm or up to several meters. The species recom- line, rocky and sandy. The three foregoing zona-
mended in rocky or coralline substrate can also tions can be generally categorized as seaward
grow well in sandy substrate. Muck substrate part of the mangroves. It was important to note
has similarity with mud except that it contains a that very few seedlings have been observed in
large amount of plant debris, this means that the these zonations maybe due to usual disturbance
organic matter is higher than mud. It also tends by waves and water current.  The propagules
to be deeper starting at more than 10 centimeters used in the experiment were planted in the open
thick up to a few meters. It has similar smell space or gap in between the mature trees. The
with mud just like foul and/or rotten egg. result showed that all the propagules planted in
 In the most seaward part of the mangroves Rhizophora stylosa zone and Avicennia marina
the substrate is coralline and rocky with some zone yielded zero survival at the end of the ex-
patches of sandy areas, which favor the growth of perimental period, which lasted for one year. In
Rhizophora stylosa, Avicennia marina and Son- the Sonneratia alba zone, 23% or 7 propagules
neratia alba. In the middle mangrove, wherein remain alive out of the 30 propagules planted.
the substrate is generally muddy, the zonations With respect to the in situ soil salinity, in Rhizo-
of Rhizophora apiculata, Bruguiera gymnor- phora stylosa zone, the salinity ranges from
rhiza and Kandelia candel are naturally grow- 8.7–19.7 parts per thousand (ppt) with annual
ing. Extending to the landward most part are the mean soil salinity of 15 ppt, while in Sonneratia
zonations of Lumnitzera littorea, Heritiera litto- alba zone, the soil salinity ranges from 9–19.6
ralis, Acanthus ebracteatus, Nypa fruticans and ppt, with similar annual mean salinity of that
Acrosticum aureum. The preferred soil type and Rhizophora stylosa zone. Sonneratia alba zone
habitat location in the mangroves of each species on the other hand, had a lower salinity ranging
is presented in Table 4. from 8.3–17.4 ppt with annual mean of 14 ppt.
 The next zones following the first three most
3.4 Identified mangroves seaward vegetations are Rhizophora apiculata,
 There were about 12 major species, 5 minor Bruguiera gymnorrhiza and Kandelia candel
species, 12 mangrove associates and 3 special- zones which based on the actual observation
ized groups identified. The category and identifi- can be generally categorized as middle part of
cation were based on the criteria used by Tomlin- the mangrove forest because they are located in
son (1986). The complete list is also presented in between seaward and landward most parts of
Table 4. the mangrove where shallow out flow of fresh
water from Baler river and an inflow of saline
water from the Pacific Ocean meets during high
4. Discussion
tide. The type of soil or substrate in these zones
 The zonation in Baler is a representative of is almost the same and can be categorized as

muddy. In terms of soil salinity, Rhizohora api- pear that Kandelia candel can withstand well in
culata zone recorded an annual salinity range of association with Nypa fruticans zonation based
4–16.7 ppt with mean of 8.1 ptt, while Bruguiera on the number of survived propagules (63%).
gymnorrhiza zone has a salinity range of 3.9–11 Lumnitzera littorea on the other side formed the
ppt and Kandelia candel zone ranges from tallest canopy in the whole forest having bigger
4–11.5 ppt. Comparatively, the last two zones trunk diameter with muck type of soil substrate,
have similar annual mean salinity of 7 ppt dur- meaning that there are more plant debris than
ing the entire duration of the observation period. mud, has small percentage survival of 13%. The
With regards to percentage survival of planted dominant canopy of this species covered almost
propagules, it was interesting to note that these all the under suppressed saplings and seedlings
three zones yielded lower mortality and almost from the direct exposure to sunlight. This might
more than half of the planted propagules sur- also one of the possibilities that contribute to
vived with 67% for Rhizophora apiculata zone, the lower survival of the planted Kandelia can-
57% for Bruguiera gymnorrhiza zone and the del propagules. The soil salinity ranges from
highest was recorded in its original zone (Kande- 4–12 ppt, with annual mean salinity of 7.8 ppt.
lia candel zone) with 70% survival rate. The Heritiera littoralis zone usually formed
 In the landward most part of the mangal area vegetation in an elevated muddy and sandy soil
which comprises of Lumnitzera littorea zone, substrates or mounds believed to be created by
Heritiera littoralis, Acanthus ebractetus, Nypa mud crabs and other soil animals and organisms
fruticans and Acrostichum aureum zones showed in mangroves that the area is usually inundated
a higher mortality except for Nypa fruticans by saline water only during high tide. Higher
which yielded 63% survival rate with soil salin- mortality of the seedlings planted in this zone
ity ranging from 0.7–4 ppt, having annual mean was recorded leaving only 1 or 3% survival.
of 2.1 ppt. The presence of small creek flowing Acanthus ebracteatus zone yielded 7% survival
in the middle of this zone might be one of the rate, with soil salinity readings ranging from 0.9–
reasons of lower salinity readings. Although, 6ppt, and with an annual mean of 3.2 ppt, which
it is hard to find open space in Nypa fruticans strategically located just closer to Nypa fruticans
zone because of its natural behaviour of forming zone. The substrate is a combination of muddy to
a dense clumps or vegetation, it seemed to ap- sandy. On the other hand, Acrostichum aureum is

Fig. 3. Percentage survival of Kandelia candel in different mangrove zonations with varying levels of
soil salinity.
10 Hikobia Vol. 15, No. 2, 2008

the farthest of all zonations but also thriving near stylosa and Avicennia marina wherein the soil
small tributaries of the creek with muddy type of salinity readings were generally higher among all
substrate. This is known as the mangrove fern in other zonations. Although, this species is con-
the Philippines because it resembles leaves simi- sidered halophyte or plant that can tolerate saline
lar to tropical ferns. The record shows that 27% environment, certainly, it has minimum and maxi-
or 8 seedlings survived in this zone after the end mum tolerance limitations. Several studies which
of experiment. The percentage survival rate of supported the above-mentioned fact include:
Kandelia candel in different mangrove zonations Kameya et al. (1996), stated that the growth inhi-
with varying levels of soil salinity is presented in bition for both Kandelia candel and Rhizophora
Fig. 3. stylosa is 1.8%, which is almost equivalent to 18
ppt. Hwang (1983) stated that this species grows
in the intertidal zone along estuarian river banks
5. Conclusion
of which the tidal salinity ranges from 10–36 ppt.
 Based on the results of this study particularly Hwang and Chen (1995) stated that Kandelia
on the number of survived propagules planted in candel seedlings grows well at salinity level up
different mangroves zones, it is concluded that to 260 mM (15 ppt) with 85 mM (5 ppt) as the
the mangroves Kandelia candel follows its origi- optimum level and with 340 mM (20 ppt) salin-
nal zonation for greater survival with soil prefer- ity, the growth will be impaired and/or inhibited.
ences to muddy soil subtrate in the middle portion Wakushima et al. (1994) revealed that the stem
of mangroves which is not expose to strong tidal elongation was observed in Kandelia candel seed-
current and with an appropriate level of in situ lings were inhibited with soil salinity of 100 mM
salinity ranging from 4–11.5 ppt or annual mean (6ppt) and a salinity concentration of 600 mM
salinity of 7 ppt. The relationship of soil salinity (35 ppt) was fatal or detrimental for it. The study
and percentage survival of this species in its orig- of Hwang and Chen (2001) revealed that salinity
inal zone is presented in Fig. 4, indicating that if was the primary factor affecting the growth of
salinity is low, the percentage survival is higher Kandelia candel. Their findings stated that the
and vice versa. The same interpretation can also uptake water salinity of Kandelia candel growing
be made in Figure 3, that the survival of Kandelia in situ was 6–14 ppt based on the Na content on
candel in zonations with high salinity readings, a its tissue. Although, their findings vary from each
lower trend of survival was noted. Zero survival other because of the effects of external factors
on the most seaward zonations of Rhizophora such as precipitation, topography, underground

Fig. 4. Survival of planted Kandelia candel in its original zone with monthly in situ soil salinity level.

water and inflow of fresh water from springs, 36: 25–31.

creeks and rivers. The findings in this study were Hwang, Y. H. & Chen, S. C. 2001. Effects of am-
gathered in the actual field and may therefore monium, phosphate, and salinity on growth, gas
vary on the level of soil salinity that could be ob- exchange characteristics, and ionic contents of
tained in an experiment conducted in a controlled seedlings of mangrove Kandelia candel (L.) Druce.
or laboratory areas such as green houses or other Bot. Bull. Acad. Sinica 45: 131–139.
artificial settings. Other possible major factors Kathiresan, K. & Bingham, B. L. 2001. Biology of
that contributed to the zero survival rate at the mangroves and mangrove ecosystems. Adv. Mar.
two seaward most zones were the disturbance Biol. 40: 1–251.
caused by strong current and wave actions, which Kameya, H., Nehira, K. & Nakagoshi, N. 1996.
sometimes brought debris that washed or injured Growth of cultivated seedlings of Kandelia candel
the planted propagules. and Rhizophora stylosa. TROPICS 6 (1/2): 51–64.
 Other zonation which yielded more than 50% Melana, D. M., Atchue, J., Yao, C. E., Edwards, R.,
survival were Rhizophora apiculata zone, Nypa Malana, E. E. & Gonzales, H. I. 2000. Mangrove
fruticans and Bruguiera gymnorrhiza zones. Management Handbook. 96 pp. Dept. Env. Nat.
Therefore, it can also be recommended that Kan- Res., Manila Philippines through the Coastal
delia candel could also be planted in these zones, Resource Management Project, Cebu City, Philip-
but careful consideration with soil preferences pines.
and salinity should not be neglected.  Other ef- Primavera, J. H. 2000. Philippine Mangroves: Status,
Threats and Sustainable Development. A paper
fects of biotic and abiotic factors such as fresh
presented during the International Workshop Asia–
water inflow, tidal inundation, water pH, dispersal
Pacific Cooperation–Research for Conservation of
ability of the propagules, physical and chemi-
Mangroves. Okinawa, Japan.
cal elements present in the soil, geomorphology,
Rabinowitz, D. 1978. Early growth of mangrove seed-
competition, pests and all others factors which
lings in Panama, and a hypothesis concerning the
may possibly play important roles in the oc-
relationship of dispersal and zonation. J. Biogeogr.
currence of zonation are stongly recommended
5: 113–33.
for specific study.  It is therefore worthwhile to
Robertson, A. I. & Alongi D. M. 1991. Tropical Man-
conclude that mangrove zonation is a natural
grove Ecosystems. 329 pp. American Geophysical
pattern or guide to follow in planting or doing
Union Press, Washington.
reforestation efforts to attain higher percentage
Saha, S. & Choudhury, A. 1995. Vegetation analysis
survival with due consideration to the type of soil
of restored and natural mangrove forest in Sagar
substrate and soil salinity requirements of each Island, Sunderbans, East Coast of India. Indian J.
species. Mar. Sci. 24: 133–136.
Semeniuk, V. 1980. Mangrove zonation along an
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Accepted 5. IX. 2008