Trees (2014) 28:1105–1113
DOI 10.1007/s00468-014-1021-2
ORIGINAL PAPER
Mangrove community structure and regeneration potential
on a rapidly expanding, river delta in Java
Sukristijono Sukardjo • Daniel M. Alongi •
Yaya I. Ulumuddin
Received: 9 January 2014 / Revised: 29 March 2014 / Accepted: 18 April 2014 / Published online: 5 May 2014
Ó Springer-Verlag Berlin Heidelberg 2014
Abstract
Key message Mangroves in rapidly expanding South-
east Asian river deltas form floristically simple zones
dominated by a few highly regenerative species adapt-
able or tolerant to rapid sedimentation and extensive
river flooding.
Abstract The size class distribution, community compo-
sition and spatial structure of five representative mangrove
forests in the rapidly expanding Cimanuk river delta on
Java were determined. These deltaic forests are species-
poor (eight true mangrove species) and spatially segregated
into three distinct floristic zones: (1) a fringing, low
intertidal zone co-dominated by Avicennia marina and A.
officinalis, with less abundant Bruguiera parviflora, Rhi-
zophora apiculata, and R. mucronata; (2) a zone transi-
tional between the low and mid intertidal in which
Avicennia and Rhizophora spp. co-dominate; and (3) a mid
intertidal zone dominated by R. mucronata and R. apicu-
lata. Numerically dominated by seedlings (52,500–73,500
seedlings ha-1) and saplings (5,268–5,660 saplings ha-1),
all five forests are relatively young and actively regener-
ating. Positive correlations of tree stem diameter and tree
height with soil organic matter and P concentrations,
salinity, the soil C/N ratio, pH, and silt/clay composition
Communicated by C. Lovelock.
S. Sukardjo  Y. I. Ulumuddin
The Centre for Oceanographic Research, Indonesian Institute of
Sciences, Jl. Pasir Putih 1, Ancol Timur,
P.O.Box 4801, JKTF, Jakarta 11048, Indonesia
D. M. Alongi (&)
Australian Institute of Marine Science, PMB 3, Townsville,
QLD 4810, Australia
e-mail: d.alongi@aims.gov.au
highlight the importance of soil factors in sustaining forest
growth. The low diversity and relative structural simplicity
of these rapidly growing and regenerating forests may be
attributed to adaptation or tolerance to flooding and the
rapid sedimentation and seaward expansion of the delta.
Keywords Community structure  Diversity  Floristic
composition  Mangrove forest  Regeneration potential 
River delta
Introduction
Indonesia possesses magnificent mangrove forests that are
not only among the most extensive but also among the
most diverse on earth (Sukardjo and Alongi 2012). And
although there have been some early floristic studies of
Indonesia’s mangroves (Becking et al. 1922; de Haan
1931; van Steenis 1958), there are only a few studies
adequately describing patterns of species distribution and
abundance and their underlying causal mechanisms (Suk-
ardjo et al. 1984; Sukardjo 1995). Such information is
sparse despite the fact that mangroves represent an
important resource and source of income for coastal
inhabitants and the government. Globally, comparatively
few studies have analyzed mangrove community structure
and diversity within the equatorial zone (Chapman 1976;
Saenger 2002), defined here as ±10° latitude of the
equator.
On the island of Java, mangroves are sites of conflict
due to incompatible activities among stakeholders, espe-
cially along the northern coast of West Java Province,
where most forests are in a degraded state (Sukardjo 1984,
1989). Only a few mangrove areas on Java are in pristine
condition. Some such forests lie within the Cimanuk delta
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1106 Trees (2014) 28:1105–1113

Indramayu (10 m alt.)

1621 mm
300

Temperature (°C)
Rainfall (mm)
100 50
80 40
60 30
Java Sea 40 20
20 10
Indonesia
J F M A M J J A S O N D J

Java Sea

deltaic

Inter-deltaic

Indramayu Legend:
Prodelta clays
Barrier sand of interdistributary area (delta)
Barrier sand of interdeltaic zone
Mouth-bar sands
Interdistributary and delta front silts
Mangroves
0 3km
Sarwah & Tegalan

Fig. 1 Mangrove forest area in the Cimanuk delta complex, Ujung Indramayu, in relation to the depositional facies in the delta (Hehannusa et al.
1976). Upper insert is a climate diagram for the meteorological station at Indramayu

complex at Tiris, approximately 15 km northwest of the
city of Indramayu, although most of the Cimanuk forests
have been subjected to severe over-cutting for decades.
With no coordinated control over these forests, it is feared
that the remaining forests will decline further or be con-
verted to fishponds. The present situation is further com-
plicated by the fact that this delta is prograding rapidly as a
result of rapid sediment discharge from the deforested
catchment of the Cimanuk River (Wolanski and Spagnol
2000; Yuanita and Tingsanchhali 2008), making it difficult
to predict changes in mangrove community structure and
composition as the delta expands.
Present investigations of the mangrove forests in Tiris
are concerned with the structure and composition of veg-
etation and physical and chemical properties of the soil as
part of a larger study of the ecology of the Cimanuk delta
complex (Sukardjo1984; Sukardjo and Yamada 1992a).
This paper, the first of a series on the delta, describes the
community composition and regenerative ability of man-
groves in this rapidly expanding river delta. We examined
current forest structure and regenerative capacity of the
dominant species so that such information will offer some
insight into the composition and survival of mangroves as
the delta progrades further into the Java Sea.

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Trees (2014) 28:1105–1113 1107

Materials and methods Table 1 Edaphic (mean ± 1 SE) factors within each mangrove zone
in the Cimanuk delta
The study area Soil parameter Sites I and II Sites III and Site V
IV
The mangrove vegetation was inventoried in 2005 at five
Silt (% DW) 19.9 ± 1.0b 21.9 ± 0.9a 17.9 ± 0.9c
different sites located on the western shore of the b b
Clay (% DW) 54.7 ± 2.8 54.7 ± 2.0 64.4 ± 2.5a
Cimanuk delta complex (6°150 S, 108°180 E) at Tiris in a a
Sand (% DW) 25.5 ± 1.1 23.5 ± 1.0 17.7 ± 0.9b
West Java Province. These sites have an area of about
pH 5.9 ± 0.8a 6.5 ± 0.6a 7.1 ± 0.6a
500 ha and are among the best-developed mangroves
Soil salinity 33.2 ± 0.5a 32.2 ± 0.6a 31.3 ± 0.9b
along the Javanese coast and were chosen to be repre- b b
Organic matter 3.4 ± 0.2 3.6 ± 0.3 4.8 ± 0.2a
sentative of the mangrove communities currently occu- (%)
pying areas subjected to rapid sedimentation and flooding
C organic (%) 2.0 ± 0.3b 2.1 ± 0.1b 2.8 ± 0.2a
(Sukardjo 1984). This flat coastal plain extends seaward b a
N total (%) 0.17 ± 0.03 0.31 ± 0.03 0.20 ± 0.07b
onto prograding tidal flats that receive riverine sediments c a
Available P (ppm) 3.7 ± 0.6 39.7 ± 1.2 25.8 ± 1.8b
at a rate of 6,000 t km-2 year-1—one of the highest c a
C/N ratio 11.5 ± 0.8 16.0 ± 0.9 13.7 ± 0.8b
recorded deposition rates in the world (Wolanski and a b
NO3 (lM) 32.4 ± 1.6 10.2 ± 1.0 11.3 ± 1.0b
Spagnol 2000)—as a direct result of unsustainable land-
NH4 (lM) 179.3 ± 6.8c 324.4 ± 5.8a 236.4 ± 6.5b
use practices (Yuanita and Tingsanchhali 2008). Diurnal
a,b,c
tides fluctuate between 0.55 to 1.75 m and annual rain- Significant differences among zones with superscript a being the
fall is 1,621 mm (see climate diagram in Fig. 1 for the highest value and c the lowest. Identical superscripts between zones
indicate no significant differences (P [ 0.05)
nearest meteorological data), but tidal water supply
within the delta is greatly influenced by levees that have
been constructed to prevent erosion. These levees are to show canopy gaps. In five transects at site V, more
still breached, resulting in extensive floods throughout detailed censuses were made of seedlings and saplings in
the delta. different-sized canopy gaps.
Soils in all plots were measured concurrently for silt,
Forest measurements sand and clay, redox potential, pH, pore water salinity, and
dissolved inorganic nutrients, and total organic carbon,
Five 0.25 ha (50 m 9 50 m) permanent plots were total nitrogen and available P using methods outlined in
established within: (1) the Avicennia zone, at site I Alongi (1996). Triplicate samples to a depth of 20 cm were
(0–50 m from sea edge) and site II (50–100 m from sea taken from each plot using a stainless steel corer (7 cm
edge); (2) the transitional zone, at site III (100–150 m inside diameter). Interstitial water was obtained from the
from sea edge) and site IV (150–200 m from sea edge); resulting core hole by syringing pore water after
and (3) the Rhizophora zone, at site V (200–250 m from percolation.
sea edge) for a total of 25 plots. The Avicennia, tran-
sitional and Rhizophora zones correspond to the low Statistics
intertidal zone, the boundary between the low and mid
intertidal zones, and the mid intertidal zone, respec- The vegetation data were analyzed to calculate species
tively, using Alongi’s (1989) definition of intertidal richness (S, total number of species present; Margalef
zonation. 1958), Shannon–Weiner diversity (H’), and (J) evenness
Tree density in each plot was estimated for three age (Legendre and Legendre 2012). An Importance Value
and diameter size classes (Sukardjo 1987): seedlings Index was calculated as the sum of relative frequency,
(dbh: B2.0 cm), saplings (dbh: 2.0–9.99 cm) and adult relative density, and relative basal area of each species
(dbh: C10 cm) trees. Each 0.25 ha plot was further (Ellison et al. 2000). The method of Pool et al. (1977)
subdivided into twenty-five 10 m 9 10 m subplots in was used to calculate a Complexity Index for a 0.1 ha
which all trees with dbh [ 10 cm were measured and plot.
recorded for estimates of stem density and basal area. One-way ANOVAs were performed to determine dif-
Some Rhizophora spp. trees were very tall, so dbh was ferences in edaphic and stand characteristics between forest
measured just above the uppermost intersection of prop sites on untransformed data (if assumptions for the test
root and main stem. Tree height was measured with a were met) followed by the Holm–Sidak procedure for
Haga altimeter and their positions were mapped. Selected pairwise multiple comparisons when significant main
subplots (10 m 9 10 m) were used for spatial mapping effects were found (Sokal and Rohlf 2012). In nearly all
to produce a crown projection diagram (adult trees only) cases, data from replicate plots were pooled as preliminary

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1108 Trees (2014) 28:1105–1113

Table 2 Forest composition of trees, saplings, and seedlings, as density (D, number ha-1), dominance (basal area, BA = m2 ha-1), average
cover (AC = % cover) and importance value (I.V.) in the Cimanuk delta
Sites Species Trees (dbh [ 10 cm) Saplings (dbh 2–9.99 cm) Seedlings (dbh \ 2.0 cm)
D BA I.V. D BA I.V. D AC I.V.

I Avicennia marina 130 1.5 113.0 1,711 4.9 81.1 9,500 3.7 50.1
A. officinalis 48 0.6 32.2 600 1.9 41.4 8,500 8.9 79.3
Aegiceras corniculatum 16 0.2 10.5 195 0.7 27.0 11,500 3.7 55.5
Rhizophora apiculata 51 0.6 34.8 649 2.2 43.2 14,500 4.1 66.9
R. mucronata 126 1.4 109.6 2,446 8.1 107.3 8,500 3.9 48.2
Total 460 5.3 5,600 17.7 52,500 24.2
II A. marina 89 1.1 83.9 1,466 5.8 71.3 8,500 3.7 43.7
A. officinalis 59 0.7 31.5 560 2.0 37.9 7,500 7.9 62.6
A. corniculatum 29 0.4 15.8 280 0.9 27.3 10,000 3.7 47.7
Bruguiera parviflora 19 0.2 10.3 180 1.9 23.6 4,500 5.5 41.9
R. apiculata 57 0.7 53.5 936 2.2 52.3 14,500 4.1 62.8
R. mucronata 112 1.4 105.2 1,844 8.6 86.7 7,500 3.8 41.4
Total 427 5.4 5,268 18.8 52,500 28.6
III A. marina – – – 100 1.0 16.9 2,000 2.4 9.5
A. officinalis – – – 80 0.8 13.5 1,500 2.4 7.2
A. corniculatum – – – 20 0.01 0.2 500 2.4 2.4
B. parviflora 58 0.7 30.6 20 0.01 0.2 7,500 2.4 35.7
B. sexangula 48 0.6 25.3 530 1.8 29.7 13,000 3.4 61.9
Ceriops decandra 19 0.2 10.3 120 0.1 2.0 12,000 7.6 57.2
R. apiculata 145 1.7 125.7 3,600 9.1 151.2 16,000 4.9 76.2
R. mucronata 124 1.5 108.2 1,500 5.6 93.1 11,000 4.6 52.4
Total 470 5.5 5,270 18.0 63,000 22.9
IV A. marina – – – 100 1.0 16.5 1,000 0.4 5.7
A. officinalis – – – 120 1.1 18.9 500 0.2 2.9
A. corniculatum – – – 40 0.02 0.3 500 0.2 2.9
B. parviflora 44 0.5 32.1 560 0.5 8.3 6,500 2.5 37.2
B. sexangula 42 0.5 30.4 220 0.4 6.7 10,000 3.8 57.2
C. decandra 28 0.3 20.3 330 0.6 10.1 10,500 4.0 60.0
R. apiculata 127 1.6 112.2 3,600 8.1 128.3 14,000 5.3 80.0
R. mucronata 120 1.5 105.1 1,500 7.1 112.8 9,500 3.6 54.3
Total 454 5.6 5,560 18.9 52,500 19.8
V A. marina – – – 100 1.2 18.9 1,000 0.4 4.1
A. officinalis – – – 100 1.2 18.9 1,000 0.4 4.1
A. corniculatum – – – 40 0.02 0.3 500 0.2 2.0
B. parviflora – – – 220 0.3 5.4 3,000 2.0 12.3
B. sexangula – – – 120 0.1 1.9 16,000 6.5 65.3
C. decandra 13 0.2 10.3 20 0.01 0.2 500 2.3 35.1
R. apiculata 236 2.9 146.2 3,700 10.6 167.6 31,000 12.7 126.5
R. mucronata 205 2.5 143.5 1,560 5.5 86.8 7,000 2.9 28.6
Total 454 5.6 5,660 19.0 73,500 30

analysis showed that differences among plots within the
same location were not significant. If necessary, data are
log10-transformed. Pearson’s product moment correlation
analysis was used to relate forest and soil factors. Results
were compared at the 1 and 5 % levels of significance.
Results and discussion
Soils in all plots (Table 1) consisted of a mixture of dark
gray silt–clay with lesser amounts of sand (18–25 %)
overlying deep clay. Eight of the 12 measured soil

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Table 3 Mean Importance Value Indices of tree species and their mean density (number ha-1, in parentheses) in each zone in the Cimanuk delta
Species Avicennia zone Transitional zone Rhizophora zone
Plot I Plot II Plot III Plot IV Plot V

Avicennia marina 113.1 (130)b 83.9 (89)a – – –

A. officinalis 32.2 (37) 31.5 (34) – – –
Aegiceras corniculatum 10.5 (12) 15.8 (17) – – –
Bruguiera parviflora – 10.3 (11) 30.6 (35) 32.1 (36) –
B. sexangula – – 25.3 (39) 30.4 (34) –
Ceriops decandra – – 10.3 (12) 20.3 (23) 10.3 (10)
Rhizophora apiculata 34.8 (40) 53.5 (57) 125.7 (145)b 112.2 (127)b 146.22 (180)b
R. mucronata 109.6 (126)a 105.2 (112)b 108.2 (124)a 105.10 (120)a 143.5 (155)a
Total mean densities 345 320 345 340 345
a b
Co-dominant, dominant

Table 4 Structural characteristics of the trees, saplings and seedlings across zones in the Cimanuk delta. Rhizophora spp values are in
parentheses
Vegetation parameter Trees Saplings Seedings

Density (number ha-1) 427–470 (97.2 %) 5,268–5,660 (93.6 %) 52,500–73,500 (53.7 %)

Basal area (m2 ha-1) 5.3–5.6 (97.9 %) 17.7–19.0 (97.5 %) –
Height ± 1 SD (m) 11.6 ± 2.5 6.8 ± 0.9 0.5 ± 0.2
Number of species (S) 3–6 5–8 5–8
0
Shannon diversity (H ) 0.4–0.5 0.4–0.8 0.8–1.0
Evenness (Shannon) (J) 0.2–0.4 0.4–0.7 0.7–0.8
Complexity Index 3.2–3.8 10.3–13.7 –
Average diameter crown ± 1 SD (m) 12.8 ± 3.5 9.6 ± 1.1 1.2 ± 0.4

parameters peaked at sites III and IV. Soil pH
(range = 5.1–7.7) did not vary significantly among plots,
but salinity was significantly less at site V. Organic matter
and organic carbon content exhibited identical patterns
with highest concentrations at site V and equivalent values
at the other sites (Table 1). The C/N ratio, P concentra-
tions, and interstitial NH4? concentrations were signifi-
cantly higher at sites III and IV with equivalent values at
the other sites. Interstitial nitrate concentrations were
higher at sites I and II than at the other sites (Table 1).
Eight species were found in the five forests with all
species numerically dominated by seedlings and saplings
rather than adults (Table 2). Total densities of trees
(427–470 individuals ha-1), saplings (5,268–5,660 indi-
viduals ha-1) and seedlings (52,500–73,500 individuals
ha-1) were within a narrow range among the five study
sites (Table 2). Composition was co-dominated by Rhizo-
phora apiculata and R. mucronata with Importance Value
Indices ranging from 34.8 to 146.2 % and 105.2 to
143.6 %, respectively (Tables 3). These forests consisted
of aggregations of adult trees of both species (40–180 trees
ha-1 and 112–115 trees ha-1, respectively), with less dense
Avicennia marina and A. officinalis (89–130 trees ha-1 and
34–37 trees ha-1, respectively). The mean number of
species (S) = 5.2 ± 0.2, mean Shannon–Weiner diversity
(H0 ) = 0.98 ± 0.38, and mean evenness (J) = 0.29 ±
0.01 varied among the five sites (Table 4). Based on the
Importance Value Index (Table 3), \25 % of the tree flora
consisted of sparse populations of Aegiceras corniculatum,
Bruguiera parviflora, Bruguiera sexangula, and Ceriops
decandra. No rare species were found.
A profile diagram (Fig. 2) and crown projection diagram
(Fig. 3) of plots IV (150–200 m from sea edge) and III
(200–250 m from sea edge) show combinations of plant
associations and age and height class differences typical of
these forests (Table 5) where three distinct floristic zones
were identified. At the most seaward sites (Plots I and II,
0–100 m from sea edge) where the frequency of tidal
inundation was maximal, the canopy co-dominants were
Avicennia marina and A. officinalis, with Bruguiera par-
viflora, Rhizophora apiculata, and R. mucronata as less
abundant species. There was a transitional zone (plots III
and IV, 100–200 m from the sea edge) landwards, in which
both Avicennia and Rhizophora spp. co-dominated. The
third zone (Plot V, 200–250 m from the sea edge) was the
most landward and dominated by both Rhizophora spp.

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1110 Trees (2014) 28:1105–1113

Fig. 2 (top) Profile diagram of 10 Rm

the mangrove forest at site IV Rm Rm
Rm Rm Rm Ra

Height (m)
and (bottom) its crown
6 Rm Rm Ra
projection in the Cimanuk delta. Rm
Ra, Rhizophora apiculata; Rm, Ra Ra
Rhizophora mucronata; Am, Ra
2
Avicennia marina; Bp, Et
Ht
Bruguiera parviflora; Et, Mt
Lt
exceptional high tide; Ht, High
tide; Mt, Medium high tide; Lt,
Low tide 10
Ra Ra Rm
Ra Am
Rm Rm
Ra Ra Ra
Rm
Rm Ra Ra Ra Am
Rm Ra Rm
Ra Ra Ra Rm Rm
Rm Rm
Rm Ra Rm
Rm Rm
Rm Rm Ra
Rm Rm Ra Ra Ra Rm
Bp Rm
Rm Am Ra
Ra
Rm 30 Bp
0 10 20 40 50 (m)

An extended relative growth relationship equation was

(m) 50
employed for determining the relationship between trunk
diameter (D, cm) and height (H, m):


40 1=H ¼ 1= aDh þ 1=Hmax
where a is a constant, h is the relative growth power index,
30
and Hmax is the maximum tree height; a, H and Hmax were
determined by weighted least-squares regression, and their
20 accuracy was examined by fitting to the C–D rule
(Shinozaki and Kira 1961): (A) 1/H = 1/(0.495D1.90) ? 1/
10 13.555 for R. apiculata; (B) 1/H = 1/(1.449D1.60) ? 1/
9.035 for mixed stand; (C) 1/H = 1/(0.958D1.35) ? 1/
0 8.225 for A. marina; and (D) 1/H = 1/(2.897D1.0) ? 1/
11.975 for R. mucronata; the relative growth power index,
h, ranged from 1.00 to 1.90.
0 10 20 30 40 50 (m)
Differences in sapling numbers between height and
Fig. 3 Crown projection diagram of the mangroves at site III in the diameter-size classes (Table 5) were likely to be caused by
Cimanuk delta. The crowns are of adult trees. Symbols: = trees canopy structure (Fig. 3) as the age class distribution
13–15 m height; = trees 12–13 m height; = trees 10–12 m shows that R. apiculata and R. mucronata saplings clearly
height; and = forest gap
dominate. The seedling composition in these stands varies

Table 5 Number of saplings and adult trees by mean diameter and height classes in all plots within the Cimanuk delta
Diameter class (cm) Height class (m) Total
1–2 2–3 3–4 4–5 5–6 6–7 7–8 8–9 9–10 10–11 11–12 12–13 13–14

2–3.99 11 21 13 3 9 4 3 2 2 – – – – 67
4–5.99 5 22 10 9 26 24 13 4 8 – – – – 121
6–7.99 3 6 7 4 38 38 9 7 6 1 – – – 99
8–9.99 – 4 1 1 16 16 4 7 17 2 – – – 66
10–11.99 – – – – – – – 6 6 13 12 14 7 57
12–13.99 – – – – – – – – 3 3 2 3 2 15
14–15.99 – – – – – – – – 2 2 3 2 9
16–17.99 – – – – – – – – – – 4 3 1 8
18–20.00 – – – – – – – – – – – 4 2 6
Total 19 53 30 27 102 75 36 27 53 21 17 23 13 586

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from 5 to 7 species with highest species diversity found in Table 6 Number of individual Rhizophora seedlings, saplings, and
plot V. Seedlings of both Rhizophora spp. represent an young trees in gap areas (n = 8) within the Rhizophora zone
important seed bank with 22,000–39,500 individuals ha-1 Gap size (m2) Number of individuals
or 46.9 ± 2.4 % of the total (Table 4) with R. apiculata
Young trees Saplings Seedlings
seedlings (importance value, I.V. = 84.9 ± 2.2) being the
dominant species. 10.8 0 1 1
In the crown projection diagram (Fig. 3), the open parts 10.8 2 1 0
(in which crowns [10 m in height were absent) were 14.5 0 2 1
regarded as gaps (blackened areas). The smallest and 15.5 0 3 7
largest size of the gaps was 10.8 and 65.8 m2, respectively. 25.6 3 2 2
The total gap area was 235.2 m2, which was 9.4 % of the 45.5 0 6 12
study area and the average gap size was 29.4 m2. The 46.5 1 6 10
forest profile and crown projection diagram of the struc- 65.8 1 8 17
tural characteristics of the forest confirm that no anthro-
pogenic disturbances such as illegal logging have recently
occurred. In the Rhizophora zone, there were a few seed- Table 7 Pearson correlation coefficients (r) of mangrove height and
lings (1–7 individuals) in the smallest gap, and denser dbh with edaphic characteristics across all plots
populations of saplings (6–8 individuals) and seedlings Soil variable Height dbh
(10–17 individuals) in the largest gaps (Table 6).
The mangrove forests of the Cimenuk delta consist of Organic matter 0.69* 0.51*
species common to other mangrove forests bordering the C/N ratio 0.17 0.18
Java Sea. These deltaic forests are species-poor compared P 0.76** 0.67**
to other mangrove patches (up to 13 species) along the pH -0.99** -0.99**
delta coast (Sukardjo and Yamada 1992b) and along the Silt/Clay 0.99** 0.99**
remainder of the northern coast of West Java Province Soil salinity 0.59** 0.53*
(Sukardjo and Alongi 2012). The lack of diversity can be * P \ 0.05 %; ** P \ 0.01 %
attributed to the very dynamic sedimentology of the
Cimanuk delta complex; the delta has been expanding into
the Java Sea since the 1960s (Purbohadiwidjojo 1964; zonation, with soil salinities being significantly less in the
Hehannusa et al. 1976). Thus, the lack of geological sta- more landward Rhizophora zone than in the two more
bility is an important variable affecting biodiversity of seaward zones. In an earlier pilot study, Sukardjo (1984)
mangroves in the delta as only species either adaptable or correlated increasing seedling height with increasing soil
tolerant to flooding and rapid sedimentation would survive salinity in the Cimanuk delta complex. This agrees well
(Sukardjo and Yamada 1992a). with Saintilan (1998) who reported that in the Mary and
The low species diversity also implies intense compe- Hawkesbury estuaries in Australia, height and dbh of
tition among and between species (Chapman 1976), with mangroves correlated positively with increasing salinity.
available light, space and nutrients likely being the main The mangroves of the Cimanuk delta receive abundant
drivers regulating forest structure and composition. Indeed, freshwater runoff and overland flow, so variations in tree
mangrove tree height and dbh correlated significantly with height (Table 4) may partly be due to the quality of the
a number of edaphic characteristics (Table 7), underscor- runoff and the frequency of daily inundation. Mangroves
ing the importance of soil texture, salinity and limiting are often nutrient-limited (Feller et al. 2002; Alongi 2009),
nutrients. A plethora of factors that are well known to play and such may be the case in the delta as mangrove height
significant roles in determining the structural characteris- and dbh correlated (Table 6) strongly with available P
tics of mangrove vegetation (Saintilan 1998; Ellison et al. concentrations (but not with total organic carbon or nitro-
2000) no doubt also operate in the delta. gen content). In contrast, growth conditions for B. par-
Three floristic zones (an Avicennia zone, a transitional viflora and B. sexangula were optimum only at sites III and
zone, a Rhizophora zone) were identified on the basis of the IV. Phosphorus limitation is possible as it occurs for these
Importance Value Index and the crown projection diagram species under laboratory (Alongi 2011) and field conditions
(Fig. 3). Populations of Avicennia marina and A. officinalis (Feller et al. 2002)
were most abundant at the seaward edge (Tables 2, 3), as The principal drivers of zonation are complex, depen-
these species are regarded as light demanding, salt tolerant dent on the interrelationships between and among factors,
pioneers. As in other mangrove forests (Ball 1988; Saint- including soil nutrients, frequency of tidal inundation,
ilan 1998), salinity appeared to be a major driver of competition and rainfall (Smith 1992). The three zones of

123
1112
mangroves in the delta are similar to those found
throughout the Indo-West Pacific region (Macnae 1968)
including in South Sumatra (Sukardjo 1987), the Malaysian
Peninsula (Watson 1928), and in Papua New Guinea
(Percival and Womersley 1975; Paijmans and Rollet 1977)
where on these wet tropical coasts, the standing crop is far
greater than those on coasts subjected to low rainfall
(Sukardjo and Alongi 2012).
Canopy gaps are important in the establishment,
recruitment and growth of mangroves (Sukardjo 1987;
Smith 1992; Sherman et al. 2000). In this area of the
Cimanuk delta, the floor of each forest is intensively
pockmarked with mounds constructed by the mud lobster,
Thalassina anomala; these crustaceans extensively biotur-
bate the soil causing oxidation and acidification (Ashton
and Macintosh 2002) but may also competitively exclude
seed predators such as sesarmid crabs (Havanods 1987;
Stieglitz et al. 2000). Seedlings are abundant in the
Cimanuk delta, growing up at 45 cm and persisting on the
low shore gradient, especially under a closed canopy. Thus,
the rates of seed predation may be low in these forests as
seedlings establish rapidly when a canopy gap opens.
Considering the age structure of these forests (Table 5), the
regeneration process seems to be carried out by trees that
are within the 5–7 m height class.
In Southeast Asia, mangrove forests with abundant
seedlings appear to coincide with mud lobsters being the
most abundant benthic organism (Abdulhadi and Suhardj-
ono 1994; Dahdouh-Guebas et al. 2000, 2002; Ashton and
Macintosh 2002) suggesting that the lobsters may facilitate
forest regeneration by excluding seed predators and
extensively bioturbating the soil. Indeed, 87, 74 and 91 %
of total mangrove densities are seedlings in forests in West
Kalimantan (Abdulhadi and Suhardjono 1994), Sarawak
(Ashton and Macintosh 2002) and in the present study
where mud lobsters are abundant.
There were very few trees (74 trees = 16.8 %) 10–14 m
in height (Table 5). Most trees (80 %) are saplings
(Table 5). The low abundance of adult trees explains the
low basal area (4.0–4.2 m2 ha-1) and correspondingly low
complexity index (3.2 ± 1.0). These forests have less basal
area as compared to mangroves in South Sumatra (Suk-
ardjo et al. 1984; Sukardjo and Alongi 2012) and in Muara
Angke-Kapuk near Jakarta (Sukardjo 1989), all with a
relatively low number of adult trees.
The relative growth power index (h) ranged from 1.0 to
1.9 indicating that these forests are relatively young and
still developing as the delta advances. There appears to be
gap enhancement of recruitment and growth in canopy
gaps as happens in mangrove forests that are thriving
(Clarke and Kerrigan 2000; Sherman et al. 2000). Rhizo-
phora seedlings appear to be successful in establishing
themselves in abundance, sprouting most readily both in
123
Trees (2014) 28:1105–1113
gaps between large trees (dbh [ 10 cm) in the transitional
zone and at higher elevation (in the case of R. apiculata)
and across the intertidal (in the case of R. mucronata),
compared with the restricted recruitment success of both
Avicennia species at the sea’s edge. This recruitment suc-
cess reinforces the notion that forest regeneration in Tiris
occurs mainly in gaps by Rhizophora and on newly formed
tidal flats by Avicennia. The change of light conditions in
gaps according to the gap size has an effect on patterns of
Rhizophora seedling regeneration (Table 6); it is advanta-
geous for young plants to drift away from the shade cast by
its parent (Smith 1992; Ewel et al. 1998; Ashton and
Macintosh 2002). Mangrove seedlings that grow from
small propagules (B1 g FW) have high mortality rates,
while seedlings that develop from large (C10 g FW)
propagules (e.g., Rhizophora) have low mortality rates.
Species with small propagules appear to establish new
cohorts annually, but quickly die (Saenger 2002).
The success of Rhizophora regeneration in gaps and of
Avicennia at the sea edge is also due in part to the differ-
ences in flooding tolerance of these species. In a study of
seedlings of six mangrove species, Youssef and Saenger
(1996) found that Rhizophora and Avicennia seedlings had
similar tolerances to waterlogging, but their tolerances
were greater than those of Bruguiera and Aegiceras.
Moreover, the Importance Value indices (Table 3) indicate
higher values for Rhizophora seedlings than for Ceriops
decandra, Aegiceras corniculatum, and Bruguiera sexan-
gula seedlings in the transitional and Rhizophora zones,
and higher values for Avicennia marina and R. mucronata
seedlings in the Avicennia zone. The seedling tolerances of
a particular species to colonize a canopy gap or a newly
formed tidal flat so rapidly and successfully is likely to be
the most important determinant of future species compo-
sition of mangroves surviving in the delta.
Acknowledgments Soil analyses were performed by the Chemistry
section of the Dept of Natural Sciences, Bogor Agricultural Univer-
sity, Bogor, Indonesia.
Conflict of interest The authors declare that they have no conflicts
of interest.
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