Fill - Form 3: Structuring of Indo-Pacic sh assemblages along

AQUATIC BIOLOGY

Aquat Biol

Vol. 5: 85–95, 2009

doi: 10.3354/ab00139

Printed March 2009

Published online February 24, 2009

INTRODUCTION

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.

Structuring of Indo-Paciﬁc ﬁsh assemblages along

the mangrove–seagrass 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 mangrove–seagrass

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

–2

. 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-

grove–seagrass 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: Mangrove–seagrass continuum · Seascapes · Connectivity · Fish assemblages ·

Trophic structuring · Juvenile habitats · Indonesia

Resale or republication not permitted without written consent of the publisher

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Aquat Biol 5: 85–95, 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 mangrove–seagrass continuum; mangrove, edge

of mangroves (the mangrove–seagrass interface), and

seagrass. The aims of this investigation were to (1) de-

termine the habitat usage of the seagrass–mangrove

continuum by ﬁsh species, (2) examine how ﬁsh assem-

blages vary with respect to trophic structure, and (3)

establish whether the seagrass–mangrove continuum

operates as habitat for juvenile coral reef ﬁsh.

MATERIALS AND METHODS

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 mangrove–seagrass

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

–2

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

Sulawesi

Wakatobi MNP

INDONESIA

I N D O N E S I A

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: 85–95, 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; www.ﬁsh-

base.org). 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 mangrove–seagrass 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 length–frequency 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 mangrove–seagrass contin-

uum. Further individual non-parametric Mann-Whitney

U-tests were conducted to determine differences in

length between each individual habitat.

RESULTS

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

–2

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

seagrass–mangrove 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 mangrove–seagrass 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 mangrove–seagrass

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

3,71

= 9.19, p < 0.001),

but no difference was found in their abundance along

the mangrove–seagrass continuum (Fig. 4). Fish and

invertebrate feeders were found in signiﬁcantly greater

numbers in the mangroves than in the seagrass (F

3,71

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

3,71

= 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: 85–95, 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).

DISCUSSION

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 mangrove–seagrass continuum.

The species assemblages within this non-estuarine

mangrove–seagrass 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 seagrass–mangrove 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: 85–95, 2009

ral variation, therefore this study cannot fully dismiss

the importance of the mangrove–seagrass 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

beneﬁted from funding from Caspian Services, Khazakstan

(SG), Glasgow NHS (SG) and Operation Wallacea (RU).

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Aquat Biol 5: 85–95, 200994

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

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