Vol.3, No.5, 331-342 (2013) Open Journal of Ecology
Relationship between the tropical seagrass bed
characteristics and the structure of the associated
fish community
Rohani Ambo-Rappe1*, Muhammad Natsir Ne ssa1, Husain Latuconsina2, Dmitry L. Lajus3
1Department of Marine Science, Faculty of Marine Science and Fisheries, Hasanuddin University, Makassar, Indonesia;
*Corresponding Author: rohani.amborappe@gmail.com
2Department of Aquatic Resources Management, Faculty of Fishery and Marine Science, Darussalam University, Ambon, Indonesia
3Department of Ichthyology and Hydrobiology, Faculty of Biology and Soil Sciences, St. Petersburg State University, St. Petersburg,
Received 24 February 2013; revised 6 May 2013; accepted 8 July 2013
Copyright © 2013 Rohani Ambo-Rappe et al. This is an open access article distributed under the Creative Commons Attribution Li-
cense, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
Structural complexity of seagrass bed including
species composition and shoot density is ar-
gued to be an important factor determining fish
assemblages. However statistical verification of
such a relatio nship is possible on ly in areas with
high species richness of seagrass and fish as-
semblages which is observed in tropical w aters.
Material for this study was collected in three
seagrass beds with different structure in Inner
Ambon Bay, Eastern Indonesia. This study pro-
vided evidence that higher structural complexity
of seagrass bed was related to the higher rich-
ness, abundance, and biomass of fish. However,
lower structural complexity of seagrass patch
should not be underestimated because it pro-
vided different habitat for various stages of life
in fish. Smaller fish preferred to occupy dense
seagrass of dominant pioneer small-sized spe-
cies (Halodule uninervis) and moved to the
lesser dense bed of climax large-sized seagrass
(Thalassia hemprichii and Enhalus acoroides)
with increasing their size. This finding is impor-
t ant for seagrass-fisheries management.
Keywords: Fish; Tropical Seagrass; Structural
Complexity; Enclosed Bay; Fis heries Management
Seagrasses are submerged aquatic plants inhabiting
marine coastal waters; they occur in the intertidal zone
and in deeper areas. They grow in beds and often form
extensive underwater meadows. Seagrasses comprise
complex communities, which include the plants and their
associated flora and fauna [1]. The presence of segrasses
enhances the marine environment by increasing the
amount of physical structure and thereby increasing the
available habitat and, consequently, increasing the abun-
dance and diversity of marine organisms [2]. Leaves and
stems of seagrasses support numerous and abundant epi-
phytes which are fed upon by small epifaunal organisms
[3], which, in turn, provide food to the fishes foraging in
the seagrass beds [1,4,5]. Fish may use seagrass for the
following purposes: temporary nursery, permanent habitat
for completion of the full life cycle, feeding area for
various life stages, and/or refuge from predation [6,7].
Seagrass beds are widely distributed in the tropical
Indo-Pacific region. They often occur adjacent to coral
reefs and mangrove forests. Overall, there are 60 des-
cribed species of seagrasses worldwide, within 12 genera,
4 families and 2 orders [8,9]. Seagrasses range from small
plants with thin leaves to large plants with thick leaves.
The order from small to large genera is the following:
Halophila < Halodule < Ruppia < Zostera/ Heterozostera
< Phyllospadix < Cymodocea < Syringodium < Amphi-
bolis < Thalassodendron < Thalassia < Enhalus < Posi-
donia [10]. Indonesian waters house about 12 species of
seagrass from 7 genera which inhabit about 30,000 km2 of
the Indonesian coastal zone. They occur in the form of
monospecific (constructed by only one species of seagrass)
or multispecific beds (constructed by two or more species
of seagrass). Indonesian seagrass meadows are generally
multispecific with up to 8 seagrass species constructed
one bed [11], however, monotypic beds of seagrasses
made up of Enhalus ac or oi des or Thalassi a hemprichii do
occur [12].
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R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342
The importance of seagrass bed as a habitat and food
source for marine animals is expected to vary with the
species composition of seagrass. Some researchers found
the differences of fish assemblages in seagrass bed repre-
sented by different species of seagrass [13-16]. Moreover,
Ambo-Rappe (unpublished work) found the abundance
and species diversity of fishes were higher in multispesific
seagrass bed with high shoot density, compared to the
monospesific seagrass bed and multispesific bed with low
shoot density. The richness of seagrass species may in-
fluence faunal assemblages because more diverse sea-
grass provide greater structural complexity and therefore
more niches for the associated plants and animals. The
physical nature of the seagrass canopy is thought to play a
major role, potentially influencing available shelter, food,
and protection from predators [17]. This fact is raising a
concern on the role of seagrass diversity on their ecolo-
gical function in marine ecosystems, in particular, because
there is a tendency of declining and/or extinction of cer-
tain species of seagrass due to climate change and other
factors [18].
Different size of seagrass beds also affect the colo-
nization of the beds by marine organisms [19-21]. The
general prediction from research on terrestrial systems is
decrease in abundance and diversity in fragmented ha-
bitats [22]. This is due to the increased negative impacts,
such as radiation, wind, and water movement that act
across smaller patches rather than in larger ones. More-
over, large patches provide greater interior areas, decrea-
sing edge impact which is mostly associated with in-
creased in predation [23]. Conversely, studies in sea-
grass systems suggest that many small seagrass patches
with higher perimeter-area ratio may increase the overall
probability of encounter by larvae or other immigrants,
thereby increasing overall colonization of the patch com-
pared to larger patches [24,25]. Moreover, the large
amount of edges associated with patchy seagrass beds
may facilitate penetration of water and food to the interior
part of seagrass patches [26]. Whereas, significantly
greater total number of infaunal macroinvertebrate taxa
was found in samples from large rather than small patches
of seagrass [20,27]. Therefore, there is no consistency of
the results on the effect of patch size on the abundance of
resident fauna. The effect of habitat size on associated
fauna may be different for different species and is highly
site- and taxon-specific [28].
Few studies have been investigated how abundance and
structure of assemblages of seagrass fauna vary among
different species of seagrass [29]. This information is par-
ticularly important for tropical area, where many species
of seagrass occur together in one meadow. The structural
complexicity of the meadow involving different seagrass
species is also need to be considered.
Effect of seagrass bed characteristics composing of
different species of seagrass on fish communities is not
easy to analyze because species richness is generally not
very large. Low number of species causes difficulties of
obtaining statistically significant effects. Because of that,
study involving many species, i.e. carried in tropical eco-
systems, are of special interest.
The objective of this study was to analyze relationship
between characteristics of seagrass bed such as species
composition, shoot density, and patch area, and the stru-
cture of associated fish community in the tropical seagrass
2.1. Study Site
This study was conducted in March - May 2011 in the
inner Ambon Bay, Maluku Province, eastern Indonesia
(Figure 1). Ambon Bay consists of two parts (inner and
outer) separated by a narrow sand bar with 12-m depth,
and characterized by different hydrological conditions.
The outer bay is deeper (up to 800 m down the slopes)
with high coral cover and connect directly to the open
Banda sea, while the inner bay is shallower (a maximum
depth of 40 m) and mainly fringed by seagrass and
mangrove (personal communication).
The inner Ambon Bay has an area of 11.72 km2 and
18.30 km of coastal line. It is a semi enclosed estuarine
bay and previously well known for its support for live-
bait fisheries for supplying skipjack fisheries [30].
This bay is characterized by tropical monsoonal climate,
which has dry season (December-February), transitional I
(March-May), rainy season (June-August), and transi-
tional II (September-November). Water temperature and
salinity fluctuate seasonally with water temperature in the
range of 24.5˚C - 31.0˚C and the salinity of 27.0 - 33.3.
Relatively small amount of water discharges into the bay
from rivers correspond to the small contribution of the
rivers in the fluctuation of temperature and salinity in this
bay (personal communicatio n).
This inner bay hosts a multispesific seagrass composed
TanjungTir am
Banda Sea
MalukuIsl and
Figure 1. Study site in the inner Ambon Bay, eastern Indonesia.
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R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342 333
of four species, namely Enhalus acoroides, Thalassia
hemprichii, Halodule uninervis, and Halophila ovalis.
The former three species of seagrass produce long strap-
like leaves, whereas the latter has oval-shaped leaves.
Both leaf length and width differ between species. E.
acoroides leaves are the biggest with 300 - 2000 mm long
and 12 - 20 mm wide, followed by T. he mprichii (length;
100 - 400 mm, width; 4 - 11 mm), H. uninervis (length; 60
- 150 mm, width; 0.3 - 4 mm), and H. ovalis (length; 10 -
40 mm, width; 5 - 20 mm) [9,31].
The composition of seagrass vary from one place to
another in the bay. There were three stations selected for
this study based on number of seagrass species occurred
in a meadow, namely: 1) Tanjung Tiram (03˚39'S;
128˚12'E) located near the entrance of the bay and has
approximately 200 m × 130 m seagrass area comprising of
four species of seagrass, E. acoroides, T. hem p ri chii , H.
uninervis, and H. ovalis; 2) Lateri (03˚38'S; 128˚13'E)
located further inside the bay and has approximately 200
m × 70 m seagrass bed consists of two species of seagrass
E. acoroides and T. hemprichii; 3) Waiheru (03˚37'S;
128˚12'E) located opposite to the second station and
consists of an approximately 200 m × 60 m monotypic
seagrass patch, E. acoroides. The distance between the
stations is approximately 2 - 3 km.
Oceanographic parameters such as depth, temperature,
salinity, pH, dissolved oxygen measured during the study
showed that intra-site variation notably exceeds variation
among sites and range from 1.0 to 1.5 m, 28.90˚C to
31.40˚C, 30.10 to 33.30 ppt, 7.91 to 8.28, and 5.49 to 6.71
mg/l, respectively.
2.2. Estimation of Seagrass Shoot Density
Seagrass density measurement was performed in each
station on March 2011 by using a systematic sampling
method according to [32]. Three 100 m line transects were
placed perpendicular to shoreline in each station. The
distance between the line transects within each station was
25 m. Ten quadrates (1 m × 1 m each) were regularly
deployed in each line transect with the distance of 10 m
from each other. Seagrass were collected from a sub
sample of 20 cm × 20 cm within each 1 m × 1 m quadrate
and washed from sediment remains before being sepa-
rated to species based on [31]. Then the shoot density of
each seagrass species was counted. Sample of sediment
was also taken from each sub quadrate for sediment grain
size analysis. Sediment samples were dry-sieved using
standard laboratory test sieves of mesh sizes 2.0 mm, 1
mm, 0.5 mm, 0,25 mm, 0.125 mm, and 0.063 mm.
2.3. Fish Sampling
Fish sampling was conducted once a month at mid-low
tide with a beach seine (1.5 m wide, 15 m long, and 500
µm mesh) on each seagrass meadow. The beach seine was
dropped in the water and manually dragged 100 m over
the seagrass bed. Two parallel beach seining were per-
formed at each station and each sampling occassion in or-
der to cover the whole seagrass meadow and reduce sam-
pling bias of the fish. All fish collected were counted,
identified to species based on standard methods [33-35],
and measured for weight (to the nearest 0.01 g) and total
length (to the nearest 0.1 cm).
There were only five species found in a wide range of
individual length sizes, namely Siganus canaliculatus,
Aeoliscus strigatus, Syngnathoides biaculeatus, Acrei-
chthys tomentosus, and Paracentropogon longispinis.
Large individuals of these species (S. canaliculatus; 16.0 -
28.3 cm, A. strigatus; 12.0 - 16.8 cm, S. biaculeatus;
19.0 - 28.3 cm, A. tomentosus; 8.0 - 10.5 cm, and P.
longispinis; 8.0 - 9.7 cm) were also analyzed for their
gonad maturity. Then, all individual fish were grouped
into juveniles and adults according to length at first
maturation available in literature [33,35] and from own
analysis of gonad.
2.4. Data Analysis
Univariate data analyses (ANOVA) were used to
analyse differences in species richness, abundance of
individual fish, and fish biomass (g wet-W) among the
three stations with different characteristics of seagrass bed.
A Bonferroni post hoc test was used for comparison of
treatment means when an F-test indicated significant
(p-value < 0.05). Before performing ANOVA, all data
were tested for normality and homogeneity of variances.
Spatial similarity in fish species composition among
stations was additionally analyzed using presence or
absence of species at each station. Juveniles and adults of
five dominant species were analyzed separately to reveal
age-specific patterns of species distribution among sta-
Shoot density of seagrass decreased in the row Tanjung
Tiram-Lateri-Waiheru (Table 1). Seagrass bed area in
Tanjung Tiram were also wider compared to other two
stations. Sediment characteristics varied among locations
and Tanjung Tiram had higher content of coarse, medium,
and fine sand. Waiheru and Lateri, on the other hand, had
higher content of very fine sediment and clay (Figure 2).
A total of 9189 individual fish representing 95 species
from 38 families were collected at the three stations. Due
to the lack of differences in the fish species richness,
abundance and biomass between months (one-way
ANOVA, p > 0.05), data obtained in different months
were pooled, and further analysis was only done on the
spatial differences of these parameters between the three
distinct seagrass beds.
Five species, which had the widest range of individual
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R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342
Copyright © 2013 SciRes.
Table 1. Shoot density of seagrass species in each station.
Shoot density (number of shoot/m 2; mean ± SE, n
= 30)
Stations Enhalus acoroides Thalassia hemprichii Halodule uninervis Halophila ovalis
Tanjung Tiram 20.67 ± 1.77 16.89 ± 2.33 56.89 ± 13.70 4.89 ± 0.89
Lateri 13.40 ± 1.31 11.00 ± 1.62
Waiheru 9.00 ± 0.28
Tanjung TiramLateriWaiheru
Percentage of sediment grain siz
Very coarse sand (1-2 mm)Coarse sand (0.5 mm)
Medium sand (0.25 mm)Fine sand (0,125 mm)
Very Fine Sand (0.063 mm)Clay (< 0,063 mm)
TanjungTiram LateriWaiheru
Numberoffishspe cies
Tan j ung Tiram LateriWaiheru
Tan j ung Tiram LateriWaiheru
Figure 2. Sediment composition at three stations.
length sizes (Siganus canaliculatus, Aeoliscus strigatus,
Syngnathoides biaculeatus, Acreichthys tomentosus, and
Paracentropogon longispinis) had also higher number of
individuals compared to the others and found in all three
stations, with exception of P. longispinis which was not
found in Waiheru station (Table 2).
These five species accounted for 70% of the total
abundance: S. canali c ul atus (50.1%), A. strigatus (9.9%),
S. biaculeatus (5.0%), A. tomentosus (3.4%), and P.
longispinis (2.3%). The result of gonad analyses showed
that individuals of the five dominant fishes had mature
gonad (and thus considered adult) at minimum sizes of
17.2, 12.7, 20.0, 8.7, and 8.5 cm length, respectively.
Based on information from gonad analyses and avai-
lable literatures on the first maturation size of fish, a high
proportion (89%) of all fish collected in this study was
categorized as juvenile. Each of the dominant species
classified into juvenile and adult (Table 3 ) showed that
Lateri and Waiheru had higher percentage of adult fish
compared to Tanjung Tiram. Almost 66% of A. strigatus
in adult stage were found at Lateri, while S. canaliculat us
and S. biaculeatus were found in 86% and 70%, res-
pectively, as adults in Waiheru.
Figure 3. Mean number of fish species, fish abundance and
biomass (mean ± SE, n = 6).
31.6% of fish species common in the three stations,
whereas similarity in species composition between sta-
tions as follow: Tanjung Tiram-Lateri (43.2%), Tanjung
Tiram-Waiheru (40.0%), and Lateri-Waiheru (42.1%) (see
also Table 2)
The number of species and abundance were signi-
ficantly different among stations (ANOVA: number of
species; F = 4.569, p < 0.05, abundance; F = 3.714, p <
0.05). Tanjung Tiram had significantly higher species
richness and abundance of individual fish than Waiheru.
However, no significant differences were found after the
Bonferroni test between Tanjung Tiram and Lateri, and
also between Lateri and Waiheru. Fish biomass was
significantly lower in Waiheru compared to the other two
stations (F = 9.420, p < 0.01) (Figure 3). There were
Seagrass habitats are associated with shallow waters
and often reported to have high abundance of juvenile
fishes [36-39]. This habitat therefore is referred to as
nursery habitat which may increase the probability of
juvenile’s survival through the provision of food and
shelter. Increased food is thought to increase growth rates,
which in turn facilitates lower mortality. The structural
R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342 335
Table 2. Fish species, number of individual, and length of fish collected in each station.
Name of family and species Tanjung
Tiram WaiheruLateri
Total number
of individuals Length (cm)Adult size
Apogon sp. + 1 3.5 5)*
Apogon fragilis (Smith, 1961) + 2 1.9 & 2.5 5)*
Apogon hoevenii (Bleeker, 1854) + + + 16 3.5 - 5.5 5)*
Apogon melas (Bleeker, 1848) + 1 7.6 10)**
Cheilodipterus quinquelineatus (Cuvier, 1828) + + 17 1.60 - 5.7 12)*
Fowleria variegata (Valenciennes, 1832) + 2 3.5 & 4.0 8)*
Balistoides viridescens (Bloch and Schneider, 1801) + 2 4.0 & 5.4 60)*
Petroscirtes mitratus (Rüppell, 1830) + + + 5 4.3 - 7.5 8)**
Petroscirtes variabilis (Cantor, 1850) + + + 29 2.7 - 9.0 12)**
Bothus pantherinus (Rüppell, 1830) + + 7 6.2 - 17.5 24)*
Engyprosopon grandisquama (Temminck & Schlegel, 1846) + 1 9.4 13)*
Pardachirus pavoninus (Lacepède, 1802) + 1 5.7 25)*
Callionymus sp + + + 20 2.5 - 5.8 12)*
Dactylopus dactylopus (Valenciennes, 1837) + + 6 5.0 - 18.0 30)*
Caranx sexfasciatus (Quoy & Gaimard, 1825) + + 13 4.5 - 12.9 42)**
Trachinotus blochii (Lacepède, 1801) + 1 15.0 58)*
Carangoides uii (Waklya, 1924) + + 5 5.6 - 11.5 25)*
Gnathanodon speciosus (Forsskal, 1775) + + 3 4.3 - 7.0 100)**
Caesio caerulaurea (Lacepède, 1801) + 1 4.5 35)*
Aeoliscus strigatus (Günther, 1860) + + + 911 3.0 - 16.8 14)*
Parachaetodon ocellatus (Cuvier, 1831) + + 5 5.4 - 8.3 18)*
Heniochus acuminatus (Linnaeus, 1758) + + 9 3.0 - 8.3 20)*
Paraplagusia bilineata (Bloch, 1787) + 1 7.0 25)*
Stolephorus indicus (van Hasselt, 1823) + 27 3.3 - 4.7 12)**
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R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342
Dactyloptena orientalis (Cuvier, 1829) + + 5 8.5 - 24 38)*
Fistularia petimba (Lacepède, 1803) + + + 53 11.5 - 49.0 185)*
Gerres oyena (Forsskal, 1775) + + + 12 9.0 - 20.3 25)*
Acentrogobius sp + + 4 6.7 - 9.5 15)**
Amblygobius phalaena (Valenciennes, 1837) + 2 1.5 & 5.8 15)*
Exyrias belissimus (Smith, 1959) + 8 8.7 - 13.6 15**
Yongeichthys nebulosus (Forskal, 1775) + 1 10.5 18)*
Istigobius decoratus (Herre, 1927) + + 21 4.9 - 7.0 12)**
Diagramma labiosum (Macleay, 1883) + 1 10.0 90)*
Choerodon anchorago (Bloch, 1791) + + 4 4.6 - 6.3 38)*
Halichoeres argus (Bloch and Schneider, 1801) + 5 5.1 - 7.0 11)*
Halichoeres chloropterus (Bloch, 1791) + + 5 10.8 - 16.1 19)*
Halichoeres melanurus (Bleeker, 1851) + + + 81 4.5 - 8.3 12)**
Halichoeres scapularis (Bennett, 1831) + + + 10 7.3 - 14.4 20)*
Halichoeres schwartzii (Bleeker, 1849) + + 40 5.7 - 10.0 12)*
Cheilinus chlorourus (Bloch, 1791) + + 3 4.8 - 12.2 45)*
Stethojulis interrupta (Bleeker, 1851) + 3 7.0 - 9.0 13)*
Lethrinus harak (Forsskal, 1775) + + + 202 3.0 - 7.6 40)**
Lethrinus lentjan (Lacepède, 1802) + 2 5.3 & 6.8 40)*
Lethrinus ornatus (Valenciennes, 1830) + + 75 3.0 - 5.7 45)**
Lethrinus variegatus (Valenciennes, 1830) + + + 146 3.8 - 11.6 20)**
Lethrinus sp + 1 5 60)*
Gazza minuta (Bloch, 1797) + 1 4.0 14)*
Lutjanus biguttatus (Valenciennes, 1830) + + 170 3.0 - 8.2 20)*
Lutjanus fulviflamma (Forsskal, 1775) + + + 9 3.7 - 14.3 25)*
Lutjanus fulvus (Forster, 1801) + + 2 4.0 & 4.9 30)*
Lutjanus lutjanus (Bloch, 1790) + 1 5.0 30)*
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Acreichthys tomentosus (Linnaeus, 1758) + + + 312 2.7 - 10.5 12)*
Mulloidichthys vanicolensis (Valenciennes, 1831) + + 2 8.0 & 8.1 24)*
Parupeneus barberinus (Lacepède, 1801) + + + 322 4.7 - 16.5 30)*
Parupeneus indicus (Shaw, 1803) + 1 9.0 40)*
Upeneus tragula (Richardson, 1846) + + + 29 5.0 - 13.3 30)**
Gymnothorax richardsoni (Bleeker, 1852) + 4 18.7 - 21 30)*
Pentapodus trivittatus (Bloch, 1791) + + + 175 2.8 - 15.3 22)**
Scolopsis ciliata (Lacepède, 1802) + + + 170 3.5 - 11.4 16)**
Lactoria cornuta (Linnaeus, 1758) + + + 18 1.8 - 13.7 46)*
Inogocia sp + 2 5.8 & 16.7 35)**
Platycephalus indicus (Linnaeus, 1758) + + + 26 7.1 - 21.8 45)**
Plotosus anguillaris (Bloch, 1794) + + 3 14.0 - 17.1 25)*
Pomacentrus tripunctatus (Cuvier, 1830) + 3 7.0 - 7.6 7.5)**
Scarus sp + + + 170 3.0 - 9.6 28)*
Leptoscarus vaigiensis (Quoy and Gaimard, 1824) + + 2 2.5 & 3.7 35)*
Centrogenys vaigiensis (Quoy & Gaimard, 1824) + 1 8.0 15)*
Cephalopholis boenack (Bloch, 1790) + 1 22.0 22)*
Epinephelus coioides (Hamilton, 1822) + + 3 5.5 - 24 44)*
Epinephelus maculatus (Bloch, 1790) + 1 14.4 35)*
Phyllichthys punctatus (McCulloch, 1916) + 1 9.5 24)*
Corythoichthys intestinalis (Ramsay, 1881) + 2 14.2 & 15 16)*
Doryrhamphus dactyliophorus (Bleeker, 1853) + 1 16.5 18)*
Hippocampus kuda (Bleeker, 1852) + + + 13 4.5 - 14.8 30)*
Syngnathoides biaculeatus (Bloch, 1785) + + + 464 8.0 - 28.3 20)*
Inimicus didactylus (Pallas, 1769) + + 5 11.5 - 13.4 18)*
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Copyright © 2013 SciRes. OPEN ACCESS
Paracentropogon longispi nis (Cuvier, 1829) + + 210 4.0 - 9.7 13)**
Pterois volitans (Linnaeus, 1758) + 1 10.7 38)*
Scorpaenopsis sp + + 20 4.6 - 8.8 18)*
Scorpaenopsis venosa (Cuvier, 1829) + + + 15 1.0 - 13.5 18)*
Synanceja horrida (Linnaeus, 1766) + + 2 7.5 & 10.0 47)*
Siganus argenteus (Quoy and Gaimard, 1825) + 1 4.5 20)**
Siganus canaliculatus (Park, 1797) + + + 4603 2.3 - 28.3 18)*
Siganus doliatus (Cuvier, 1830) + + 3 2.6 - 7.5 20)*
Siganus lineatus (Linnaeus, 1835) + 1 3.8 30)*
Siganus punctatus (Schneider, 1801) + + + 32 2.2 - 5.8 24)*
Sphyraena pinguis (Günther, 1874) + 4 8.7 - 13.5 35)**
Saurida gracilis (Quoy & Gaimard, 1824) + + + 135 3.7 - 17.5 28)**
Saurida tumbil (Bloch, 1795) + 1 15.6 43)*
Pelates quadrilineatus (Bloch, 1790) + + 193 3.0 - 10.8 20)**
Arothron immaculatus (Bloch and Schneider, 1801) + + 5 2.5 - 20.7 30)*
Arothron manilensis (Marion de Procé, 1822) + + + 51 1.8 - 19.5 31)*
Arothron reticularis (Bloch and Schneider, 1801) + + + 45 2.2 - 25.0 30)*
Arothron stellatus (Bloch and Schneider, 1801) + + + 3 16.5 - 34.4 90)*
Chelonodon patoca (Hamilton, 1822) + + + 72 1.5 - 14.2 20)*
Note: (+) Found, () Not Found, )* = Allen (1999), )** = Kuiter & Tonozuka (2001).
Table 3. Percentage of juvenile and adult in five dominant species at each station.
Tanjung Ti ram Lateri Waiheru
Dominant Species
Juvenile Adult Juvenile Adult Juvenile Adult
Siganus canaliculatus 99.9 0.1 98.0 2.0 14.0 86.0
Aeoliscus strigatus 43.6 56.4 34.2 65.8 100.0 0.0
Syngnathoides biacule atus 69.5 30.5 55.7 44.3 30.0 70.0
Acreichthys tomentosus 89.1 10.9 64.1 35.9 67.5 32.5
Paracentropogon longispi nis 51.4 48.6 52.2 47.8 0.0 0.0
complexity of seagrass habitats is also considered to
provide shelter from predators [40].
In this study, approximately 89% of fish were found to
be juveniles. This fact may suggest that seagrass in inner
Ambon Bay act as a nursery habitat for the fishes around
that area. However, care should be taken for this con-
clusion as in [41] suggested examination of several
factors beside juvenile density, such as juvenile survival,
R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342 339
growth, and movement to adult habitats in order to con-
firm whether a habitat is a nursery. In most cases, how-
ever, higher density of juveniles in coastal water allows
to interpret them as nurseries. Moura et al. [42] found
juvenile of a reef fish dog snapper (Lutjanus jocu) (size <
7 cm) associated with estuarine habitat and moved off-
shore with increasing size and then considered the estua-
ry as a nursery ground of this fish. Seagrass and man-
grove were also considered as nursery habitats for some
coral reef fishes after finding higher densities of juve-
niles in these habitats and observing the pattern of mi-
gration of the fish to the coral reefs at increased size
The composition of seagrass species affected the asso-
ciated fish communities in this study. Station with more
species of seagrass (Tanjung Tiram, 4 species; Lateri, 2
species) had more fish species, and also higher abundance
and biomass of fish compared to station with one segrass
species (Waiheru). However, numbers of seagrass species
occurring in a bed might not be a solely factor contributed
to the result. Seagrass bed in Tanjung Tiram and Lateri
were also denser, and especially Tanjung Tiram had larger
area of seagrass beds as well. Structural complexity (in
this case measured as shoot density and number of
seagrass species) which was different among stations is
considered as a major factor responsible for fish richness.
Horinouchi [45] suggested that within-patch structural
complexity provides a refuge against predators, attenu-
ation in strong water movements and varied micro-
habitats, and allows for the coexistence of potentially
competing species, thereby supporting high species
richness. Horinouchi and Sano [46] found the abundance
of juveniles of three gobiid fishes positively increased
with the leaf height and shoot density of seagrass Zostera
marina. Moreover, Hemminga and Duarte [1] added that
high density of seagrass increase the surface area for
attachment of microscopic animals and plants (epiphytes)
which is the main food for fish.
On the other hand, our study reports an interesting
finding when fishes of larger size (up to adult) are more
abundant in stations which composed of only one or two
species of seagrass with less shoot density (Lateri and
Waiheru). Seagrass bed in Lateri was composed of the
climax Indo-Pacific seagrass species, Thalassia hem-
prichii and Enhalus acoroides, and Waiheru was only
composed of E . acoroides. We suggested that there would
be a movement of fish from Tanjung Tiram (which com-
posed of 4 species of seagrass, but dominated by Halodule
uninervis) to Lateri and Waiheru at increased size of fish.
Interestingly, this effect can be facilitated by migration
patterns on the earlier stages of the life cycle because the
Tanjung Tiram is closer to the entrance of that bay and
thus may early accept young fish arriving from the open
sea. This finding concurred with other studies, for
example, Middleton et al. [47] which found smaller fish
species and individuals dominated in Zostera beds,
whereas larger species and individuals were found in
Posidonia. Juveniles of several species were thought to
move from Zostera (middle size seagrass) to Posidonia
(big size seagrass) with increased size. Kendrick and
Hyndes [48] also observed the migration of spotted
pipefish Stigmatopora argus from the narrow-leaved
Posidonia coriacea to the broad-leaves P. sinuosa. A
similar trend was also found in Atlantic croaker (Micro-
pogonias undulatus) and red drum (Sciaenops ocellatus)
which smaller sizes occurred in small seagrass (Halodule
wrightii) and the bigger ones were in a big seagrass
(Thalassia testudinum) [49]. The reason for that move-
ment could be different, Kendrick and Hyndes [48]
proposed that the movement of the fish to the broader leaf
size with increased size of the fish to enable better
camouflaged to prevent predation, and Rooker et al. [49]
suggested that fish may select habitat where mortality is
Morphology of seagrass species in term of leaf size
(length and width) could be considered as an important
factor determining habitat selection by fish at each of their
life stage. In this study, beside those parameters, spacing
among seagrass shoot could result in the migration of the
bigger size of fish. Adult fish due to their bigger size will
select bigger space to enable them to move within the
seagrass bed and still protecting them from predator by
choosing bigger size of seagrass (longer and wider leaf)
which have bigger canopy. This result is in line with [15]
which stated that fish would select the habitat that match
their size showing from their study where bigger size of
fish occurred in open space below the canopy of Am-
phibolis griffithii, while smaller fish able to penetrate and
occupied the dense foliage of Posidonia sinuosa. Stoner
[50] tested the protective ability of seagrasses and found
that for single seagrass species, predation intensity de-
clined with plant surface area, however multi-species
seagrass which possessing greatest amount of total sur-
face area provided least amount of protection. The reason
for this is that spacing in the dense multi-specific seagrass
bed does not match the size of the prey that being attacked
by visual predators.
The role of physical factors (e.g. waves) could also be
considered in framework of this study as in [29] suggested
that wave actions could cover and uncover seagrass patch
as seagrass leaves sway back and forth with wave passage,
and seagrass may thus provide less protection from
predator. In this study, Tanjung Tiram is located next to
the mouth of the bay; as a result, wave action in this place
is more intense compared to other two stations which
located further inside the bay. The result of the stronger
wave action could be seen from the sediment grain size
Copyright © 2013 SciRes. OPEN ACCESS
R. Ambo-Rappe et al. / Open Journal of Ecology 3 (2013) 331-342
(Figure 2) that showed Tanjung Tiram has larger per-
centage of big grain size (sand) compared to other two
stations. It could be assumed that protection capacity of
seagrass bed in Tanjung Tiram is suitable only for smaller
size fish (or juvenile) and became weaker with increasing
size of the fish due to the many factors described above.
This pattern is also in agreement with a hypothetical
model proposed by [51], in which, seagrass bed in
Tanjung Tiram due to its close proximity to entrance
channel will be firstly encountered by fish larvae leads to
increased number of species and individuals in this station.
After settling or their size is large enough, individuals
redistribute to select the cover of other seagrass bed that
favors survival. Distance from the bay entrance with
combination with the restricted water circulation within
the bay may also limits larva dispersal, and leads to li-
mited juvenile recruitment to seagrass beds further away
from the entrance. This finding is similar to [52].
It can be concluded that the specific role of seagrass
bed for fish could be determined by its structural com-
plexity (measured by shoot density and seagrass species
composition), seagrass surface area, and physical para-
meters (such as wave and current). All the parameters
should be considered together and the effect would be
different due to factor interaction for different fish species
and their life stage. Smaller patches and less dense of
seagrass should not be overlooked because our result
demonstrated that these seagrass patches play another role
for bigger size and adult fishes. Therefore our study
reveals that different life stages of fish may find optimal
condition in different seagrass habitats. It means that it is
not some particular seagrass habitat is most important for
fish community than others, but presence of different
seagrass habitats, i.e. their heterogeneity is the most
important factor for maintaining fish populations. The
role of seagrass meadow as a habitat of fish is well known,
however, more study still need to be done to understand
many specific questions regarding to this role because it
could be site- and species-specific. Identifying what kind
of seagrass habitat is used for different fish species and at
each life stage of fish is crucial for seagrass-fisheries
We would like to thank all colleagues who assisted in the field and lab
works. Funding for field expenses was provided by COREMAP II and
Darussalam University to H.L. Manuscript preparation was partly
funded by Indonesian Higher Education (PAR-C) to R.A-R to visit St.
Petersburg State University, Russia.
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