Aquat Biol
Vol. 5: 8595, 2009
doi: 10.3354/ab00139
Printed March 2009
Published online February 24, 2009
Mangroves often occur in close association with
coral reefs and seagrass beds, forming a complex
ecosystem linked by biological and physical processes
(Parrish 1989, Dorenbosch et al. 2005, Sheaves 2005,
Unsworth et al. 2008). One third of the world’s man-
grove forests have been lost in the past 50 yr, mostly
due to human activities (Alongi 2002), and are declin-
ing at rates possibly faster than coral reefs and tropical
rainforests (Duke et al. 2007). Similarly, high rates of
seagrass loss have also been observed throughout the
world (Orth et al. 2006). Understanding the degree of
utilisation of these habitats by fish communities is
important in aiding the development and implementa-
tion of effective resource management programs.
© Inter-Research 2009 ·*Email:
Structuring of Indo-Pacific fish assemblages along
the mangroveseagrass continuum
Richard K. F. Unsworth
1, 5,
, Samantha L. Garrard
, Pelayo Salinas De León
Leanne C. Cullen
, David J. Smith
, Katherine A. Sloman
, James J. Bell
Northern Fisheries Centre, Department of Primary Industries and Fisheries, PO Box 5396, Cairns, Queensland 4870, Australia
School of Biological Sciences, University of Plymouth, Drake Circus, Plymouth PL4 8AA, UK
Centre for Marine Environmental and Economic Research, School of Biological Sciences, Victoria University of Wellington,
PO Box 600, Wellington, New Zealand
CSIRO Sustainable Ecosystems, James Cook University, PO Box 12139, Earlville BC, Cairns, Queensland 4870, Australia
Coral Reef Research Unit, Department of Biological Sciences, University of Essex, Wivenhoe Park, Colchester CO4 3SQ, UK
ABSTRACT: Indo-Pacific mangrove swamps and seagrass beds are commonly located in close prox-
imity to each other, often creating complex ecosystems linked by biological and physical processes.
Although they are thought to provide important nursery habitats for fish, only limited information
exists about their usage by fish outside of estuaries. The present study investigated fish assemblages
in non-estuarine intertidal habitats where mangroves and seagrass overlap (the mangroveseagrass
continuum). Three habitats (mangrove, mangrove edge, seagrass) were sampled at 4 sites of the
Wakatobi Marine National Park, Indonesia, using underwater visual census. Ninety-one species of
fish were observed at a mean density of 130.1 ± 37.2 ind. 1000 m
. Predatory fish (fish that feed on
invertebrates and/or fish) were the most dominant feeding groups in the mangroves, whilst omni-
vores dominated on the mangrove edge and in the seagrass. Although the habitats along the man-
groveseagrass continuum were observed to be important for many fish, only 22 of the 942 coral reef
species known within the area utilised mangroves as nursery habitat and only 15 utilised seagrass.
Despite finding evidence that nursery grounds in mangroves and seagrass may not directly support
high coral reef fish diversity, many of the coral reef nursery species found in this study are likely to
be key herbivores or apex predators as adult fish on local coral reefs, and thus highly important to
local fisheries. Although mangroves are not permanently inundated by the tide, this study highlights
their importance as fish habitats, which at high tide support a greater abundance of fish than seagrass
beds. In the light of the high rate of destruction of these habitats, their role in supporting fish assem-
blages requires consideration in marine resource management programs.
KEY WORDS: Mangroveseagrass continuum · Seascapes · Connectivity · Fish assemblages ·
Trophic structuring · Juvenile habitats · Indonesia
Resale or republication not permitted without written consent of the publisher
Aquat Biol 5: 8595, 2009
The role of non-estuarine mangroves as a nursery
ground for coral reef fish remains highly controversial
(Nagelkerken 2007, Nagelkerken et al. 2008). Although,
within regions such as tropical Australia, estuarine
mangroves have commonly been found to harbour
high densities of juvenile fish (Robertson & Duke 1987,
Blaber et al. 1989, Blaber et al. 1995) and make a large
contribution to coastal fisheries productivity, these are
not always coral reef fish. Within the Indo-Pacific,
mangroves have often been considered to play a very
limited role as nursery habitats for coral reef fish (Par-
rish 1989, Williams 1991, Laroche et al. 1997, Nagelk-
erken et al. 2008, Unsworth et al. 2008). The manage-
ment of coral reefs throughout the Indo-Pacific is of
critical importance due to their continuing degradation
(Wilkinson 2008). Knowledge of nursery habitats asso-
ciated with these coral reefs is important in making
informed management decisions.
Several reasons have been proposed for the use of
mangroves and seagrass beds by fish as juvenile habi-
tats, including: (1) their function as a refuge from pre-
dation (Parrish 1989, Robertson & Blaber 1992), (2) the
abundance of feeding resources (Hemminga & Duarte
2000, Baldo & Drake 2002), (3) their ability to intercept
planktonic fish larvae (Parrish 1989), (4) the reduced
predator density (Parrish 1989), and (5) the turbidity
decreasing the foraging efficiency of predators (Robert-
son & Blaber 1992). These roles have largely been
investigated in studies of highly turbid estuarine man-
groves with various ecological processes and in clearer
water mangroves that do not have interactions with
nutrient-rich freshwater flows.
Seagrass and mangrove habitats are best known for
their ‘nursery’ function. However, many species of fish
spend their whole lifecycle in these shallow water habi-
tats (Sheaves 2005), whilst many adult fish routinely
undertake tidal and diel migrations between these and
adjacent habitats (Nagelkerken et al. 2000, Dorenbosch
et al. 2004, Nagelkerken & van der Velde 2004,
Unsworth et al. 2007a,b). In the Indo-Pacific, tidal
ranges are often high and mangroves can commonly be
alternately flooded or exposed during the tidal cycle
(Sheaves 2005). Apart from some species from the fam-
ily Gobiidae, which have specialised air-breathing ca-
pabilities (Park et al. 2006), fish that utilise mangroves
during high tide must migrate to adjacent lower habi-
tats when the tide is low (Vance et al. 1996). These mi-
grations away from mangroves at low tide are likely to
be towards neighbouring deeper habitats such as sea-
grass beds (Vance et al. 1996, Marguillier et al. 1997,
Lugendo et al. 2006), suggesting that the biological
connectivity between these 2 habitats is prominent.
As well as possibly supporting the productivity of
coral reef fish communities, Indo-Pacific non-estuarine
mangroves may also harbour high densities of addi-
tional fish species. Many species of fish are thought
to be common to both seagrass and mangrove habitats
(Baelde 1990, Yáñez-Arancibia et al. 1993, Unsworth
et al 2008), whilst seagrass beds close to mangroves
show greater fish diversity than seagrass beds alone
(Nagelkerken et al. 2001, Lugendo et al. 2005,
Unsworth et al. 2008). This enhanced diversity appears
to show a dependence on a combination of mangroves
and seagrass beds, showing that an interaction
between the two habitats confers an advantage to fish,
enhancing species richness (Unsworth et al. 2008).
The present study investigated fish assemblages of
the mangroveseagrass continuum; mangrove, edge
of mangroves (the mangroveseagrass interface), and
seagrass. The aims of this investigation were to (1) de-
termine the habitat usage of the seagrassmangrove
continuum by fish species, (2) examine how fish assem-
blages vary with respect to trophic structure, and (3)
establish whether the seagrassmangrove continuum
operates as habitat for juvenile coral reef fish.
Study Design. The present study was conducted dur-
ing July and August 2006 in the Kaledupan sub-region
of the Wakatobi Marine National Park (WMNP), a
group of islands off SE Sulawesi, Indonesia, known for
their high diversity of marine life (Halford 2003). This
coastal zone is characterized by high coverage of sea-
grass, mangroves and coral reefs. Seagrass beds con-
sisted of mixed seagrasses dominated by Thallassia
hemprichii and Enhalus acoroides (Unsworth et al.
2007c). Mean percentage seagrass coverage ranged
from 48.8 to 79.5% (Unsworth et al. 2008). Mangrove
forests were dominated by Rhizophora spp., although
lower densities of Avennicia spp. and Brugiera spp.
were found at both Darawa sites. Semi-diurnal tides in
this area show an average tidal range of approximately
1.5 m (Unsworth et al 2007a), with intertidal mangrove
forests exposed at low tide.
Four sites were chosen (Fig. 1); Darawa Village
(05°55’54.2’S, 123°86’ 40.4’E), Darawa Mangroves
(05° 54’ 80.6‘’ S, 123° 87’12.1’E), Languira (05° 47’
57.9’S, 123° 71’91.0’E) and Sombano (05° 48’02.1’ S,
123° 69’ 86.4’E). The 4 sites are all adjacent to small
offshore islands and are exclusively marine, with no
riverine influence. They border onto seagrass mead-
ows and lie in close proximity to coral reefs. They are
at a distance of least 100 m from the land and located
well away from the immediate vicinity of a large popu-
lation centre; consequently, they were not considered
to be affected by anthropogenic factors.
The only freshwater input is from diffuse terrestrial
run-off during the wet season. Environmental char-
Unsworth et al.: Structuring of Indo-Pacific shallow-water fish assemblages
acteristics of the individual sites are summarized in
Table 1. The mangrove sites are located near reef envi-
ronments and adjacent to seagrass habitats; seagrass
habitats are within a deeper intertidal range and not as
abundantly exposed as mangroves. Further descrip-
tions of these sites can be found in Unsworth et al.
(2008). To study the abundance and size distribution of
different fish species along the mangroveseagrass
continuum, 3 habitats were distinguished: mangrove
(inside the mangroves; at least 50 m into the mangrove
and away from the outer edge, and not within large
open mangrove channels), mangrove edge (the very
outer seaward facing edge of the mangrove), and sea-
grass (50 m away from the mangrove outer seaward
edge in continuous seagrass cover).
At each of the 4 sites, 6 repeat visual transects were
conducted within each of the 3 habitats yielding a total
of 72 transects. In order to conduct the visual transects,
observers were trained at estimating fish lengths under-
water by repeatedly estimating the lengths of objects
of known length underwater (see English et al 1997).
Habitats were sampled using an underwater visual
census (UVC) method with snorkelling equipment and
50 m belt transects. Transects in the seagrass were laid
50 m into the seagrass parallel to the mangroves, and
transects on the edge of the mangroves were laid 3 m
from the edge and parallel to the mangroves. In these
2 habitats, fish were counted in an area 2.5 m either
side of the transect tape, giving a total sampling area of
250 m
. Due to the shade and decreased visibility
within the mangroves, fish were counted within 1 m
either side of the transect, giving a total area of 100 m
In order to compare data between habitats and with
other studies, all data is presented as ind. 1000 m
Prior to laying transects, a number of preliminary site
visits were conducted in order to determine suitable
places to conduct transects, i.e. those places where an
observer could move between mangrove trees and
prop roots at high tide. Conducting these mangrove
belt transects was extremely challenging due to the
dense forest. For example, observers commonly
pushed their way through tight branches in order to
continue transects. Unfortunately, some transects could
not be fully completed and were instead broken and
restarted at the nearest possible location. Although not
a commonly used methodology, mangrove UVC stud-
ies have previously been conducted in Rhizophora
mangrove of both the Indo-Pacific and the Caribbean
(Nagelkerken & van der Velde 2002, Dorenbosch et al.
2005, 2007, Aguilar-Perera and Appeldoorn 2007). Dif-
ficulties associated with the use of UVC within man-
groves, such as observer bias and fish behaviour, are
extensively discussed by Nagelkerken & van der Velde
(2002). All transects were conducted during daylight
between 07:00 h and 17:00 h at periods of high tide,
when the mangroves and seagrass beds were fully
flooded. On a couple of occasions, conditions between
mangroves were too turbid and dark to conduct obser-
vations, particularly in locations where sand had a
dark colour; therefore these transects were conducted
at the next available occasion. As with any visual
observation method in any vegetated environment, it
is likely that small and cryptic species were underesti-
mated (Edgar et al. 2004). Many mangroves in and
around estuarine environments are commonly highly
turbid, shallow, and inappropriate for UVC of fish
assemblages. However, the present study was based
Wakatobi MNP
0 400 km
Fig. 1. Location of the 4 study sites within the Kaledupa sub-region of the Wakatobi Marine National Park, SE Sulawesi, Indonesia
Table 1. Environmental and location characteristics (mean ± SE) of
the 4 sites located in the Kaledupa sub-region of Wakatobi Marine
National Park, Indonesia. All measurements were taken along the
seaward edge of the mangrove
Site Visibility Temp Salinity Distance
(m) (°C) (‰) to coral
reef (km)
Langeira Beach 5.3 ± 0.2 28.4 ± 0.0 34 ± 2 0.6
Sombano Beach 4.7 ± 1.0 30.4 ± 0.0 33 ± 3 0.9
Darawa Mangroves 4.5 ± 0.1 28.3 ± 0.0 34 ± 2 0.9
Darawa Village 6.0 ± 0.1 28.3 ± 0.0 33 ± 2 1.2
Aquat Biol 5: 8595, 2009
within non-estuarine mangroves surrounding small off-
shore islands. At high tide, mangroves in the WMNP
become inundated by clear oceanic waters that allow
for high visibility and water depths commonly exceed-
ing 1.5 m. Despite the clear water, the observation of
fish species within shaded prop roots remained diffi-
cult and required a number of training sessions prior to
observations being undertaken (Nagelkerken & van
der Velde 2002).
Data analysis. All summary statistics are presented as
means ± SE, with the exception of size distribution
across habitats, which was presented as median values
± interquartiles due to data following a Poisson distrib-
ution. Fish species were classified as juveniles when
under 1/3 their maximum length unless their maximum
length exceeded 90 cm, in which case they were classi-
fied as juveniles when under 30 cm (after Nagelkerken
& van der Velde 2002). Maximum lengths were based
on data in Fishbase (Froese & Pauly 2008; Each fish species was assigned a feeding cat-
egory (fish and invertebrate feeder, invertivore, herbi-
vore, omnivore or planktivore) after Unsworth et al.
(2007a). Multiple sources were used in order to ensure
placement of species in the correct trophic feeding cat-
egory (Hutomo & Peristiwady 1996, Khalaf & Kochzius
2002, Nakamura et al. 2003, Froese & Pauly 2008).
All univariate analyses were performed using
MINITAB 15 software. Data was tested for model confor-
mity to meet assumptions for analysis of variance
(ANOVA) using Levene’s test and, where necessary, the
data was log (x + 1) transformed to meet the require-
ments. Comparisons of overall fish abundance, species
richness, abundance of each trophic category and juve-
nile and adult abundances between sites and habitats
along the mangroveseagrass continuum were analyzed
using 2-way ANOVAs. Post-hoc Tukey’s tests were con-
ducted to determine individual inter-site differences.
Multivariate analysis of community assemblage data was
conducted using Primer v6 software (Plymouth Marine
Laboratory). The Bray-Curtis similarity index was ap-
plied to square-root transformed data (to reduce the
influence of abundant and rare species), resulting in a
triangular similarity matrix. A non-metric multidimen-
sional scaling (nMDS) ordination of similarity matrices
was used to test for significant differences in species
assemblages between habitats at each site. Confirmation
of differences in assemblage structure was conducted
using analyses of similarities (ANOSIM). A similarity
percentage (SIMPER) analysis was used to determine
which species were contributing to the differences ob-
served by the ANOSIM (Clarke & Warwick 1994). It was
not possible to use parametric tests to analyze size dis-
tribution between habitats, as data followed a Poisson
distribution and did not conform to the assumptions of
ANOVA. Comparisons of lengthfrequency distribu-
tions between habitats using two-sample Kolmogorov-
Smirnov tests showed that distributions were signifi-
cantly different between habitats. A Kruskall-Wallis test
was used to perform the non-parametric equivalent of an
ANOVA, looking at the difference in fish lengths be-
tween habitats along the mangroveseagrass contin-
uum. Further individual non-parametric Mann-Whitney
U-tests were conducted to determine differences in
length between each individual habitat.
Fish assemblage composition
A total of 91 species from 32 families were observed
during the study period (Appendix 1, Table A1). The
average fish abundance was 130.1 ± 37.2 ind. 1000 m
and the average fish species richness was 6.3 ± 0.8 spe-
cies. Numerically, the most dominant families were:
Atherinidae (55.5%), Apogonidae (19.2%), Siganidae
(5.9%) and Labridae (3.4%). In terms of numbers of
species per family, the most dominant were: Labridae
(11 species, 12.1%), Pomacentridae (10 species,
11.0%), Apogonidae (6 species, 7.0%) and Nemipteri-
dae (6 species, 7.0%).
Total fish abundance differed significantly along the
seagrassmangrove continuum (Table 2). Post-hoc
Tukey’s tests showed that fish abundance was signifi-
cantly greater in the mangroves than on the mangrove
edge or in the seagrass (Fig. 2a). No difference was
Table 2. Two-way GLM ANOVA results for adult, juvenile and total fish abundances in 3 habitat types (mangrove, mangrove
edge and seagrass) across 4 sites in the Kaledupa subregion of the Wakatobi Marine National Park, Indonesia. Fish were sampled
using underwater visual census. Seq SS: sequential sum of squares
Juveniles Adults All Fish
Source df Seq SS F p Seq SS F p Seq SS F p
Site 3 1.0 0.9 >0.1 1.6 2.0 0.119 1.2 2.5 >0.5
Habitat 2 10.4 13.7 < 0.0001 20.2 39.0 <0.0001 27.2 82.2 <0.0001
Interaction 6 0.5 0.2 >0.1 2.5 1.6 0.17 0.7 0.7 >0.1
Error 60 22.8 15.6 9.9
Total 71 34.8 39.9 39.1
Unsworth et al.: Structuring of Indo-Pacific shallow-water fish assemblages
found in fish abundances between study sites. Species
richness showed significant variability between sites
= 11.17, p < 0.001), but not between habitats along
the mangroveseagrass continuum (Fig. 2b). Post-hoc
Tukey comparisons showed that the Languira site
showed significantly greater species richness than the
other study sites (p < 0.01). Richness did not vary
between the other study sites.
Fish assemblages varied significantly (Global R =
0.619, p < 0.01) throughout the mangroveseagrass
continuum (Fig. 3). No significant difference in assem-
blage composition was found between sites. Atheri-
nomorus lacunosus and Lutjanus ehrenbergii were the
most representative species of the mangrove habitat,
Scolopsis trilineatus and Choerodon anchorago were
most representative of the mangrove edge, and Lethri-
nus harak and Siganus canaliculatus were most repre-
sentative of the seagrass habitat (Table 3).
Trophic structuring
There was a significant difference in the abundance
of omnivores between sites (F
= 9.19, p < 0.001),
but no difference was found in their abundance along
the mangroveseagrass continuum (Fig. 4). Fish and
invertebrate feeders were found in significantly greater
numbers in the mangroves than in the seagrass (F
4.38, p = 0.017), although there was between-site vari-
ability within this trophic category. Invertivores did not
show between-site variability and were consistently
found in higher abundances in the mangroves than in
the seagrass (F
= 4.32, p = 0.018). The herbivore
Fig. 2. (a) Abundance and (b) richness of fish assemblages
(mean ± SE) within the mangrove, mangrove edge and sea-
grass habitat sites sampled. Different letters indicate signifi-
cant differences between habitats as determined by post-hoc
Tukey comparisons (p < 0.01)
Fig. 3. nMDS plot of fish assemblages in different habitats
within 4 different sites of the Kaledupa sub-region of the
Wakatobi Marine National Park. Fish assemblages were sam-
pled using underwater visual census. V: Darawa Village, D:
Darawa Mangroves, L: Languira; S: Sombano
Table 3. SIMPER analysis of individual habitat similarity to determine those fish groups, in terms of abundance, that comprise
>4% similarity within all 3 habitats (mangrove, mangrove edge and seagrass). SIMPER was calculated using fish abundance data
recorded using underwater visual census at 4 sites (Darawa, Darawa Village, Languira and Sombano) in the Wakatobi Marine
National Park. % SC: percent similarity contribution
Mangrove Mangrove edge Seagrass
Fish species % SC Fish species % SC Fish species % SC
Atherinomorus lacunosus 47.8 Scolopsis trilineatus 22.2 Lethrinus harak 30.0
Lutjanus ehrenbergii 9.1 Choerodon anchorago 17.7 Siganus canaliculatus 27.7
Lethrinus harak 6.4 Gerres oyena 10.8 Gerres oyena 9.6
Sphaeramia orbicularis 5.0 Lethrinus harak 9.8 Scolopsis trilineatus 7.2
Scolopsis ghanam 4.9 Acanthurus grammoptilus 7.1 Siganus fuscescens 6.5
Scolopsis trilineatus 4.8 Siganus canaliculatus 6.9 Halichoeres argus 4.8
Dischistodus fasciatus 5.7
Goby sp. 5.1
Lutjanus ehrenbergii 4.2
Aquat Biol 5: 8595, 2009
feeding category contained the fewest individuals
(Fig. 4). These herbivores were found in highest abun-
dances along the mangrove edge, although variability
was significant between sites as well as between habi-
tats. Atherinomorus lacunosus and Sphaeramia orbicu-
laris had to be removed from the analysis; due to their
shoaling nature, their very high abundances skewed
any comparative analysis. A. lacunosus were the
only planktivorous fish which occurred at these sites
throughout this study period. S. orbicularis are fish and
invertebrate feeders. Both species were found almost
exclusively within the inner mangrove habitat.
Juvenile and length distributions across habitats
There were no significant differences in juvenile or
adult abundances between study sites; however, the
abundance of both juveniles and adults was signifi-
cantly higher within mangroves than in the other 2
habitats (Fig. 5, Table 2). Fish size differed signifi-
cantly overall between habitats (χ
= 1646.520, p <
0.001) (Mann-Whitney U-test); however, median fish
length (Fig. 6) was greater in seagrass beds than in any
of the mangrove habitats where little difference
between habitats was observed.
Of the 91 species recorded within seagrass and man-
grove habitats, 50 have previously been recorded on
local coral reefs (Halford 2003). Mangrove habitats
were found to contain juveniles of 39 species, at least 22
of which have previously been recorded as adult fish on
local coral reefs (Halford 2003). Of the 26 species found
as juveniles within seagrass habitat, 15 have previously
been recorded on local coral reefs (Halford 2003).
The results of the present study show that non-estu-
arine mangroves and seagrass beds are important
habitats for diverse fish assemblages within the Indo-
Pacific and that they also have a nursery function.
However, this function may be limited for coral reef
species. Many of the abundant species (e.g. Lethrinus
harak) were found to inhabit all 3 habitats (mangrove,
mangrove edge, seagrass), indicating that individuals
may move between them. An important finding of the
present study was that distinct assemblages were pre-
sent within all 3 habitats irrespective of site, indicating
Fig. 4. Abundances of fish in different feeding categories in
each of the 3 habitats: (mangrove, mangrove edge, seagrass)
within the Kaledupa sub-region of the Wakatobi Marine Na-
tional Park. Results are means ± SE for each habitat across the
4 study sites. Different letters indicate significant (p < 0.05)
differences between habitats
Fig. 5. Abundance of adults and juveniles (mean ± SE) in 3
habitats (mangrove, mangrove edge, seagrass) within the
Kaledupa sub-region of the Wakatobi Marine National Park.
Comparisons of juvenile and adult abundances were made
using 2-way GLM ANOVA. Significant (p < 0.05) Tukeys pair-
wise differences between juvenile fish abundance in each
habitat are represented by letters; adults by numbers
Fig. 6. Median sizes of fish (± interquartile ranges) found in
mangrove, mangrove edge and seagrass habitats within the
Kaledupa sub-region of the Wakatobi Marine National Park.
Fish sizes were surveyed using underwater visual census
Unsworth et al.: Structuring of Indo-Pacific shallow-water fish assemblages
that many species may inhabit specific zones within
this mangroveseagrass continuum.
The species assemblages within this non-estuarine
mangroveseagrass continuum consisted of many spe-
cies found in comparable earlier studies in this region
(Thollot 1992, Laroche et al. 1997, Dorenbosch et al.
2005). Mangrove habitats were numerically dominated
by species of the family Atherinidae, which is consis-
tent with the findings of studies throughout northern
Australian estuarine mangroves (Robertson & Duke
1990, Laegdsgaard & Johnson 2001). The high density
of Atherinomorous lacunosus likely indicates an abun-
dant source of plankton within these mangrove forests.
Other families of fish commonly found within man-
groves in the present study (e.g. Lethrinidae, Lutjanidae,
Gerridae) contain many species found in mangroves of
other studies (Thollot 1992, Laroche et al. 1997).
Although there were many similarities with other
studies, the fish assemblages found within mangroves
in the present study were strikingly different from other
Indo-Pacific mangrove studies due to a high density of
the family Apogonidae. Abundant species of Apogo-
nidae, such as Sphaeramia orbicularis and Apogon ce-
ramensis, are predators of small fish and crustaceans.
The specific reasons for these differences are not clear;
however, the abundance of particular crustaceans or
greater visibility may make hunting by species of
Apogonidae within these non-estuarine mangroves
more successful than in other estuarine mangroves that
have been studied throughout this region (Quinn &
Kojis 1985, Sasekumar et al. 1992, Thollot 1992).
Many studies have demonstrated that the presence
of mangroves increases species density in adjacent
seagrass beds (Nagelkerken et al. 2001, Lugendo et al.
2005, Jelbart et al. 2007, Unsworth et al. 2008), sug-
gesting that the presence of mangroves confers some
form of advantage to fish. Of the 91 species found
within seagrass and mangroves in the present study,
50 have previously been recorded in reef habitats. It is
likely that, in the present study, the flooded mangroves
are only a temporary fish habitat, and refuge is sought
in adjacent habitats such as seagrass and reef when
the tide goes out. This illustrates that many species
utilise both mangrove and seagrass habitats, and that
some coral reef and seagrass fish will utilise the shelter
and resources of a nearby mangrove habitat (Nagel-
kerken et al. 2008).
Fish that fed either wholly or partly on a diet of inver-
tebrates contributed to the majority of species present in
mangroves. This is consistent with earlier studies, which
found little evidence for fish species utilising mangrove
productivity directly. However, species of Mugilidae,
which were frequently observed during the present
study, can consume significant amounts of mangrove
detritus (Sasekumar et al. 1992), and both Siganidae and
Pomacentridae may utilise mangrove flora as food (Mar-
guillier et al 1997). The diet of carnivorous fish varies
throughout their life cycle, with an increase in prey size
with increasing length (Baldo & Drake 2002), which is
likely to reduce competition. Juveniles of large pisci-
vorous fish such as Sphyraenidae and Carangidae were
observed foraging around the mangrove/seagrass in-
terface, probably looking for and chasing schools of
fish such as hardyheads (R. K. F. Unsworth pers. obs.).
Such predatory fish may be important in structuring
fish assemblages within these mangroves. Species of
Sphyraenidae and Carangidae are more likely to use
physical structure to increase predation efficiency, rather
than as protection (Verweij et al. 2006).
The distinct assemblages present within each habi-
tat suggests that, although the shelter of these man-
groves is likely to confer advantages for some species
of fish, this is not true for all. Refuge from predation is
likely to be of greater importance to smaller fish within
these habitats. The median size of fish in the man-
groves and on the mangrove edge was much smaller
than in the seagrass. This is likely due to the habitat
complexity (Nagelkerken & van der Velde 2002) and
shade (Verweij et al. 2006) offered by the mangroves,
which are likely to reduce predation pressure and
influence smaller fish to preferentially utilise the man-
grove habitat. Larger fish, which are less at risk from
predation, may prefer to remain in the seagrass beds
where food is abundant (Ogden & Zeimann 1977).
The abundance of juveniles observed during this
study was low in comparison to the abundance of
adults, which is in contrast to recent studies in other
non-estuarine mangroves (Aguilar-Perera & Appel-
doorn 2007). Lugendo et al. (2005) suggested that
seagrass beds within the Indo-Pacific may function as
corridors between mangroves and coral reefs for reef
fish that undertake ontogenic migrations, from shallow-
water habitats during the juvenile phase to the reef,
once they mature. The present study found evidence
that only a small number of the 942 species of fish ob-
served on nearby coral reefs may utilise seagrass as
such a corridor. Only 22 coral reef species utilised man-
grove and only 15 utilised seagrass as nursery habitats.
Although we found evidence that mangrove and sea-
grass may not directly support high coral reef fish diver-
sity, it is important to realise that, of the juvenile fish
that were recorded within the seagrassmangrove con-
tinuum, some individual species are prominent apex
predators and herbivores as adults on nearby coral
reefs (e.g. Caranx ignoblis and Naso vlamingii) and are
hence likely to be important in structuring reef commu-
nities (Mumby et al. 2007). Spawning in many fish fam-
ilies varies temporally (Victor 1986, Leis 1993). The pre-
sent study, like many previous studies in the tropics,
lacks year-round data and thus information on tempo-
Aquat Biol 5: 8595, 2009
ral variation, therefore this study cannot fully dismiss
the importance of the mangroveseagrass continuum
as a ‘nursery’ area for reef fish, but indicates a need for
more year-round analysis.
The species found to utilise seagrass and mangrove
as nursery habitats in the present study, although not
hugely diverse, may also be of high ecomomic im-
portance. Many of these are critically important to
local fisheries (e.g. Siganidae, Lethrinidae, Lutjanidae,
Caranx spp., Sphyraena spp.) (May 2005, Cullen 2007);
hence, these habitats require consideration within
resource management programs.
In conclusion, seagrass beds and non-estuarine man-
groves in the Indo-Pacific support species-rich fish
assemblages. Carnivorous fish were the most domi-
nant feeding group found at high tide in the man-
groves, whilst omnivores dominated on the mangrove
edge and in seagrass. The number of herbivores was
consistently low, although it is well documented that
very few species of fish feed on mangrove or seagrass
material directly (Cebrian & Duarte 1998). Seagrass
and mangrove were found to be important habitats for
juveniles of some reef fish species. Juveniles of 22 coral
reef species were found in mangrove and 15 in sea-
grass (Halford 2003). Although mangroves are not per-
manently inundated by tides in the Indo-Pacific, this
study highlights their importance as fish habitats, sup-
porting a greater abundance than seagrass beds dur-
ing high tide. In the face of high rates of destruction
and resource extraction, their role as an important fish
habitat must be taken into consideration when devel-
oping effective resource management programs.
Acknowledgements. The authors thank all staff at Hoga
Marine Research Centre that provided logistical support
during data collection, in particular La Amat. This project
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Appendix 1. Table A1. Abundances of juveniles and adults (mean ± SE) of the fish species observed per 1000 m
in each of the three habitats
(mangrove, mangrove edge and seagrass) in the Kaledupa sub-region of the Wakatobi Marine National Park, SE Sulawesi, Indonesia.
Fish were sampled in each of the habitats using underwater visual census at high tide. Adults and juveniles of all species were categorised
by their usage of mangrove and seagrass habitats. h: herbivore; i: exclusive invertebrate feeder; f: fish-and-invertebrate feeder; o: omnivore;
p: planktivore
Taxon Juveniles Adults
Reef Trophic Man- Mangrove Seagrass Man- Mangrove Seagrass
species group grove edge grove edge
Seagrass adult and juvenile
Naso vlamingii (Valenciennes, 1835) Y h 0 3 ± 2.83 1.5 ± 0.83 0 0.33 ± 0.33 0
Dischistodus chrysopoecilus (Schlegel & Müller, 1839) h 0 0 1.5 ± 0.93 0 0 1.83 ± 1.18
Cheilio inermis (Forsskål, 1775) Y f 0 0.17 ± 0.17 0.33 ± 0.23 0 0.17 ± 0.17 0.17 ± 0.17
Gerres acinaces (Bleeker, 1854) i 0 0.17 ± 0.17 0.33 ± 0.33 0 0.67 ± 0.67 0
Rhinecanthus aculeatus (Linnaeus, 1758) Y f 0 0 0.33 ± 0.33 0 0 0
Cheilodipterus quinquelineatus (Cuvier, 1828) Y f 0 0 0.17 ± 0.17 0 0 0.67 ± 0.46
Diodon liturosus (Shaw, 1804) Y i 0 0 0.17 ± 0.17 0 0 0
Lethrinus obsoletus (Forsskål, 1775) o 0 0 0.17 ± 0.17 0 0 0
Seagrass adult only
Apogon hoevenii (Bleeker, 1854) Y i 0 0 0 0 0.5 ± 0.5 1 ± 1
Dischistodus perspicillatus (Cuvier, 1830) Y h 0 0 0 0 0.17 ± 0.17 0.67 ± 0.67
Parapercis millipunctata (Günther, 1860) o 0 0 0 0 0 0.5 ± 0.5
Amblygobious phalaena (Valenciennes, 1837) o 0 0 0 0 0 0.33 ± 0.23
Arothron manilensis (de Procé, 1822) i 0 0 0 0 0 0.33 ± 0.23
Corythoichthys haematopterus (Bleeker, 1851) i 0 0 0 0 0 0.33 ± 0.33
Leptoscarus vaigiensis (Quoy & Gaimard, 1824) Y h 0 0 0 0 0 0.33 ± 0.33
Balistapus undulatus (Park, 1797) Y f 0 0 0 0 0 0.17 ± 0.17
Chrysiptera parasema (Fowler, 1918) Y i 0 0 0 0 0 0.17 ± 0.17
Dascyllus aruanus (Linnaeus, 1758) Y o 0 0 0 0 0 0.17 ± 0.17
Halichoeres margaritaceus (Valenciennes, 1839) Y f 0 0 0 0 0 0.17 ± 0.17
Pardachirus pavoninus (Lacepède, 1802) i 0 0 0 0 0 0.17 ± 0.17
Pomacentrus lepidogenys (Fowler & Bean, 1928) Y i 0 0 0 0 0 0.17 ± 0.17
Pseudomonacanthus macrurus (Bleeker, 1857) Y o 0 0 0 0 0 0.17 ± 0.17
Syngnathoides biaculeatus (Bloch, 1785) Y i 0 0 0 0 0.67 ± 0.39 0.17 ± 0.17
Mangrove juvenile only
Labridae sp.2 f 0 1.67 ± 1.67 0 0 0 0
Labridae sp.3 f 0 0.67 ± 0.67 0 0 0 0
Siganus punctatus (Schneider & Forster, 1801) Y h 0 0.67 ± 0.67 0 0 0 0
Carangoides oblongus (Cuvier, 1833) f 0 0.33 ± 0.33 0 0 0 0
Scolopsis bilineatus (Bloch, 1793) Y f 0 0.17 ± 0.17 0 0 0 0
Mangrove adult and juvenile
Atherinomorus lacunosus (Forster, 1801) p 556 ± 481 33.3 ± 33.3 0 2261 ± 9730 83.3 ± 52.1 0
Hyporhamphus dussumieri (Valenciennes, 1847) f 108 ± 63.9 0 0 12.2 ± 9.61 0 0
Neomyxus leuciscus (Günther, 1872) o 70.6 ± 48.2 0 0 70.6 ± 28.2 5.17 ± 5.17 0
Apogon ceramensis (Bleeker, 1852) i 33.3 ± 28.9 0 0 83.3 ± 52.5 1.67 ± 1.67 0
Siganus guttatus (Bloch, 1787) Y o 10.6 ± 4.97 2.17 ± 1.32 0 2.22 ± 1.92 1.5 ± 1.17 0
Scolopsis temporalis Y i 2.22 ± 1.49 0 0 6.67 ± 4.20 0.33 ± 0.33 0
Lutjanus ehrenbergii Y f 1.67 ± 0.78 0.17 ± 0.17 0 16.7 ± 3.70 7.5 ± 2.85 0
Sphyraena barracuda (Edwards, 1771) Y f 1.67 ± 1.44 0.5 ± 0.28 0 2.22 ± 0.87 0 0
Dischistodus pseudochrysopoecilus (Allen & Robertson, 1974)
Y h 1.11 ± 0.96 0 0 2.22 ± 1.92 0 0
Caranx ignobilis (Forsskål, 1775) Y f 1.11 ± 0.96 0.83 ± 0.83 0 0 0 0
Lutjanus russelli (Bleeker, 1849) Y f 1.11 ± 0.96 0 0 0 0 0
Terapon jarbua (Forsskål, 1775) f 0.56 ± 0.48 0.17 ± 0.17 0 6.11 ± 4.81 0.5 ± 0.5 0
Acanthurus nigrofuscus Y h 0.56 ± 0.48 0 0 1.11 ± 0.96 0.67 ± 0.67 0
Plectorhinchus lessonii (Cuvier, 1830) f 0.56 ± 0.48 0 0 1.11 ± 0.66 0 0
Acanthurus fowleri (de Beaufort, 1951) h 0.56 ± 0.48 0 0 0 0 0
Arothron stellatus (Bloch & Schneider, 1801) Y i 0.56 ± 0.48 0 0 0 0 0
Toxotes jaculatrix (Pallas, 1767) o 0.56 ± 0.48 0 0 0 0 0
Pomacentrus taeniometopon (Bleeker, 1852) h 0 0.33 ± 0.33 0 6.11 ± 5.29 0 0
Unsworth et al.: Structuring of Indo-Pacific shallow-water fish assemblages
Appendix 1 (continued)
Mangrove adult
Sphaeramia orbicularis Y f 0 0 0 956 ± 415 0 0
Abudefduf septemfasciatus (Cuvier, 1830) o 0 0 0 4.44 ± 2.55 0 0
Dischistodus fasciatus (Cuvier, 1830) h 0 0 0 4.44 ± 2.13 10.3 ± 3.27 0
Hemiglyphidodon plagiometopon (Bleeker, 1852) Y h 0 0 0 3.33 ± 2.10 0.83 ± 0.59 0
Pentapodus caninus (Cuvier, 1830) Y f 0 0 0 1.67 ± 1.05 0.5 ± 0.5 0
Lutjanus decussatus (Cuvier, 1828) Y f 0 0 0 1.11 ± 0.66 0.17 ± 0.17 0
Chaetodon lunula (Lacepède, 1802) Y o 0 0 0 0.56 ± 0.48 0 0
Siganus virgatus (Valenciennes, 1835) Y h 0 0 0 0.56 ± 0.48 0 0
Acreichthys tomentosus (Linnaeus, 1758) Y o 0 0 0 0 0.17 ± 0.17 0
Bleniidae sp. o 0 0 0 0 0.17 ± 0.17 0
Blenniela paula o000 000
Ephinephelus ongus (Bloch, 1790) Y f 0 0 0 0 0.17 ± 0.17 0
Exyrias bellisimus (Smith, 1959) i 0 0 0 0 0 0
Lutjanus fulvus (Forster, 1801) Y f 0 0 0 0 0.33 ± 0.33 0
Scarus globiceps (Valenciennes, 1840) h 0 0 0 0 5.33 ± 3.01 0
Scolopsis lineatus (Quoy & Gaimard, 1824) Y i 0 0 0 0 3 ± 2.29 0
Upeneus sundaicus (Bleeker, 1855) Y i 0 0 0 0 0.17 ± 0.17 0
Seagrass and mangrove juvenile
Liza vaigiensis (Quoy & Gaimard, 1825) o 11.1 ± 6.60 4.17 ± 2.13 0.83 ± 0.59 0 0.67 ± 0.52 0.5 ± 0.5
Parupeneus macronemua (Lacepède, 1801) i 0.56 ± 0.48 0.17 ± 0.17 0.5 ± 0.37 0 0 0.17 ± 0.17
Parupeneus barberinus (Lacepède, 1801) Y i 4.44 ± 2.35 1.83 ± 0.72 1.5 ± 0.75 0 0 0
Lethrinus ornatus (Valenciennes, 1830) Y f 0.56 ± 0.48 0.17 ± 0.17 1.5 ± 0.86 0 0 0
Caranx melampygus (Cuvier, 1833) Y f 0.56 ± 0.48 0.5 ± 0.5 0.5 ± 0.5 0 0 0
Holgymnosus doliatus (Lacepède, 1801) Y f 4.44 ± 2.73 1 ± 1 0.33 ± 0.23 0 0 0
Seagrass and mangrove adult and juvenile
Siganus canaliculatus Y o 1.11 ± 0.96 1.33 ± 0.75 0.17 ± 0.17 45 ± 12.9 26.2 ± 8.62 35 ± 12.1
Choerodon anchorago Y f 21.7 ± 7.42 11.3 ± 4.65 1 ± 0.55 29.4 ± 11.3 13.2 ± 3.23 2.33 ± 1.02
Acanthurus grammoptilus (Richardson, 1843) h 5 ± 2.13 0.83 ± 0.42 0.5 ± 0.5 13.9 ± 4.03 11.7 ± 3.22 0.17 ± 0.17
Scolopsis ghanam f 3.33 ± 0.99 1.33 ± 0.71 0.17 ± 0.17 12.2 ± 3.33 1.17 ± 0.56 0
Lethrinus harak (Forsskål, 1775) Y f 22.2 ± 6.13 10.3 ± 2.58 11.7 ± 2.71 11.7 ± 4.04 1.83 ± 1.05 0.83 ± 0.34
Siganus fuscescens (Houttuyn, 1782) Y o 3.89 ± 2.34 2 ± 0.93 0.5 ± 0.37 6.11 ± 2.72 2.67 ± 1.48 32 ± 14.9
Halichoeres trimaculatus (Quoy & Gaimard, 1834) Y f 0.56 ± 0.48 0 0.17 ± 0.17 5.56 ± 3.37 0.33 ± 0.23 0
Gerres oyena (Forsskål, 1775) o 1.11 ± 0.66 14.8 ± 6.24 2.17 ± 0.93 1.67 ± 0.78 3.17 ± 0.83 4.83 ± 1.93
Lethrinus erythropterus (Valenciennes, 1830) Y f 3.33 ± 1.71 0.67 ± 0.67 0.33 ± 0.23 0.56 ± 0.48 0.17 ± 0.17 0
Seagrass and mangrove adult, juvenile mangrove
Scolopsis trilineatus Y i 0 0.17 ± 0.17 0 40.6 ± 11.4 25.3 ± 4.69 7 ± 3.49
Seagrass and mangrove adult, juvenile seagrass
Halichoeres argus (Bloch & Schneider, 1801) f 0 0 1.17 ± 0.74 4.44 ± 2.99 1 ± 0.84 8.83 ± 3.96
Halichoeres scapularis (Bennett, 1832) i 0 0 0.67 ± 0.52 1.67 ± 1.05 2 ± 1.2 0.5 ± 0.5
Halichoeres papilionaceus (Valenciennes, 1839) i 0 0 0.5 ± 0.5 1.11 ± 0.66 0 2.83 ± 1.26
Seagrass and mangrove adult
Gobiidae sp. o 0 0 0 5 ± 1.44 4.33 ± 1.15 2.33 ± 0.99
Lethrinus variegatus (Valenciennes, 1830) Y i 0 0 0 0.56 ± 0.48 0.33 ± 0.23 1.5 ± 0.58
Apogon hartzfeldii (Bleeker, 1852) Y 0 0 0 22.2 ± 19.2 0 0.33 ± 0.33
Amblygobius sphynx (Valenciennes, 1837) i 0 0 0 2.78 ± 1.69 0.5 ± 0.37 0.33 ± 0.23
Labridae sp.1 i 0 0 0 0.56 ± 0.48 0 0.17 ± 0.17
Myrichthys colubrinus (Boddaert, 1781) f 0 0 0 0.56 ± 0.48 0 0.17 ± 0.17
Taeniura lymma (Forsskål, 1775) f 0 0 0 0.56 ± 0.48 0 0.17 ± 0.17
Total 874 ± 475 95 ± 40.5 28.7 ± 3.98 3649 ± 9830 219 ± 55.4 108 ± 24.4
Taxon Juveniles Adults
Reef Trophic Man- Mangrove Seagrass Man- Mangrove Seagrass
species group grove edge grove edge
Editorial responsibility: Hans Heinrich Janssen,
Oldendorf/Luhe, Germany
Submitted: June 16, 2008; Accepted: January 8, 2009
Proofs received from author(s): February 17, 2009