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Wetlands Ecology and Management 10: 189195, 2002.

2002 Kluwer Academic Publishers. Printed in the Netherlands.

189

Benthic fauna in tropical tidal flats a comparative perspective


Sabine Dittmann
Zentrum fr Marine Tropenkologie, Fahrenheitstr. 6, 28359 Bremen, Germany,
E-mail: sabine.dittmann@zmt.uni-bremen.de

Key words: benthic assemblages, biogeographic comparison, diversity, macrobenthos, spatial distribution, species
richness, tidal flat, tropics

Abstract
Tidal flats extend seaward from mangrove forests along many tropical coastlines. This contribution compiles the
current knowledge on tropical tidal flats with regard to species richness, abundance, spatial distribution patterns
and ecological roles taken by major organisms. Tidal flats encompass a variety of soft-sediment habitats which are
inhabited by a species-rich fauna. Species numbers are generally higher than in temperate tidal flats, but vary widely
between tropical sites where they have not yet been fully assessed due to lack of research and taxonomic problems.
The assessment of biodiversity is further complicated by a low species frequency, the small size of macroinfaunal
organisms and variations in the occurrence of species between sites. Mean abundance ranges from 10002000
individuals m2 , although figures ten times higher have been recorded in a mudflat in Costa Rica, and there
are great variations in general. Individual densities of small-sized macrofauna are about ten times higher than for
larger macrobenthos. A generalized zonation scheme for intertidal distribution patterns of tropical tidal flat fauna is
presented. The comparison shows that the occurrence of macrofauna and especially of ecosystem engineers varies
between sites, which can in turn explain the great variability in species abundance patterns recorded in benthic
surveys of tidal flats within and between regions. This review revealed similarities as well as differences between
various tropical tidal flats, which underlines the need for further comparative studies to be done, using the same
methodology, before generalizations can be made.

Introduction

Comparisons of intertidal fauna between temperate


and tropical locations produced contradictory reports
on latitudinal trends in species diversity (Alongi,
1989, 1990; Reise, 1991; Vargas, 1996) and biomass
(Piersma et al., 1993; Ricciardi and Bourget, 1999),
but confirmed lower individual densities in tropical
tidal flats. Yet, how representative are the sites used
for these comparisons? How similar are tropical tidal
flats compared to each other? With limited knowledge
on variation within the tropics, a general pattern for
latitudinal comparisons of tidal flats can not be accomplished. The purpose of this literature survey is
to look for similarities as well as differences between
tropical tidal flats in order to facilitate future global
comparisons.

Dittmann and Vargas (2001) were the first to carry


out a comparison of tidal flat fauna between two tropical regions in Northern Australia and Central America. The compared sites belong to different biogeographic areas but had a high degree of faunal similarity, although variations in species diversity between
sites were high in each region. They found both generalities and singularities in the benthic assemblages of
tidal flats on both sides of the Pacific. Here, I include
studies from Africa and Southeast Asia in the comparison to obtain a more general perspective of tropical
tidal flats.
This article compiles the current knowledge on
tropical tidal flats using certain aspects relevant for
a global comparison. Based on previous investigations on tropical tidal flats it may be hypothesized
that they are rich in species, low in abundance and
similar in community structure (Macnae and Kalk,

190
1962; Day, 1974; Warwick and Rushwayuni, 1987;
Reise, 1991; Dittmann, 1995; Michaelis and Wolff,
2001). Spatial distribution patterns in relation to intertidal gradients are known from temperate and tropical
locations (Beukema, 1976; Whitlatch, 1977; Dankers
and Beukema, 1983; Ricciardi and Bourget, 1999;
Dittmann, 2000) and I will compile a generalized
zonation scheme for tropical tidal flat fauna. As it
has been proposed that structural and functional similarities between benthic assemblages originate either
from closely related species occupying similar niches
or unrelated species performing analogue ecological
functions (Reise, 1991; Vargas, 1996), I will compare
the occurrence of burrowing species between various
tropical tidal flat locations.
In the literature surveyed, the sampling procedures
varied with regard to the size of the corer, sampling
depth and sieve size used as well as the subsequent
treatment of samples. Some sites were studied only
once while other studies were more encompassing.
The studies also varied in their approach (e.g. being quantitative or qualitative) and the presentation
of data. The comparison is restricted to macrofauna
and only records from intertidal sand- to mudflats are
compiled here, leaving out beaches, seagrass beds and
mangrove forests.

Species richness
Diversity increases towards the tropics in many marine habitats (Thorson, 1957; Sanders, 1968; Ormond
et al., 1997). Does this trend apply to tidal flats as
well? Alongi (1989, 1990) and Vargas (1996) deny an
increase in diversity towards the tropics, while other
studies revealed higher species numbers at lower latitudes (Macnae and Kalk, 1962; Day, 1974; Reise,
1991; Dittmann, 1995; Michaelis and Wolff, 2001).
Table 1 compiles records of species numbers from
temperate and tropical tidal flats. In general, species
richness is higher in the tropical locations, but species numbers vary greatly within the tropics with some
sites having lower values comparable to temperate
locations and several other sites yielding very high
records. Thereby both high and low species numbers
occur within biogeographic regions.
However, this comparison remains tentative until
the species stock of tropical tidal flats has been fully
assessed. At most sites only selected taxa were treated
to species level because of taxonomic problems. Surveys are often incomplete and species-area curves in-

dicate that records will increase with further sampling


(Reise, 1991; Dittmann, 1995). Furthermore, the use
of smaller sieve sizes can increase species records,
since small sized polychaetes are especially rich in
species in tropical tidal flats (Dittmann, 1995, 2000,
in press). Species assessment is further complicated
by a low species frequency. Available records show
that between 10 and 32% of the species account for 80
or 95% of the individuals (Vargas, 1987; Wolff et al.,
1993; Dittmann, 1995; Pepping et al., 1999). Thus,
while few species tend to be abundant, the majority of
the species is represented by very few individuals.
Species compositions also tend to be site-specific
in the tropics. Even abundant and prominent macrobenthic species of one area were absent when compared to another tidal flat about 200 km away (northwestern Australia: Pepping et al., 1999; Piersma et al.,
unpubl.; north-eastern Australia: Dittmann, 2000, in
press). This restricted occurrence of species complicates generalizations and reveals the need for the use
of several sites for global comparisons.

Abundances
Lower abundances in tropical versus temperate tidal
flats were shown in a comparison by Reise (1991).
Yet, how great is the variation within tropical tidal
flats? Mean values from various tropical locations
range from around 10002000 indivduals m2 for
macrofauna, despite far higher records from a mudflat in Costa Rica, and variations are high (Table 2).
The use of smaller sieve sizes increases individual
densities. For example, 712 times as many mesothan macrofaunal organisms were found at two sites
in north-eastern Australia (Table 2, Dittmann, 1995).
Still, compared to several hundred thousand individuals m2 in temperate tidal flats, the lower values from
tropical locations compiled here confirm the finding of
Reise (1991).
Based on quantitative benthic surveys, the rank order of abundance of macrobenthic organisms can be
compared between several tropical tidal flats. Table 3
presents the five most abundant species from tidal flats
in West Africa, Australia and Central America and reveals a great variation among benthic assemblages, as
each tidal flat has a distinct set of common taxa.

191
Table 1. Total numbers of macrozoobenthic species reported from tidal sand- and mudflats around the world. Sandy
beaches or sites within mangrove forests were not considered for this comparison. Methods differed in the cited
studies and mesh sizes of the sieves used are provided.
Location

Species number

Method used, mesh size

Temperate tidal flats


Wadden Sea, Germany 1)
Barnstable Harbour, Massachussetts, USA 2)
Tomales Bay, California, USA 3)
Bahia Quillaipe, Chile 4)

78
82
74
43

1 mm
0.75 mm
1.5 mm
0.25, 0.5, 1 mm

Tropical tidal flats


Banc dArguin, Mauretania 1)
Inhaca Island, Mozambique 5)
Morrumbene estuary, Mozambique 6)
Ao Nam Bor, Thailand 4)
Selangor, Malaysia 7)
Tanjong Tokong, Malaysia 8)
Mata Ikan, Singapore 9)
Bay of Rest, West Australia 10)
Anna Plains, West Australia 11)
Roebuck Bay, West Australia 12)
Hinchinbrook Channel, Queensland, Australia 13)
Haughton River estuary, Queensland, Australia 14)
Estero de Jaltepeque, El Salvador 15)
Punta Morales, Costa Rica 16)
Ilha de Canela, Brazil 17)

132
465
292
118
51
92
140
112
114
161
202
96
76
108
77

0.6 mm
Handpicking, sieving
Handpicking, sieving (1 mm)
0.25, 0.5 mm
Handpicking
Handpicking
Handpicking, sieving (1, 1.5, 5 mm)
1, 2 mm
1 mm
1 mm
0.25, 1 mm
0.25, 0.5, 1 mm
0.5 mm
0.5 mm
0.5 mm

1) Michaelis and Wolff (2001), 2) Sanders et al. (1962), 3) Johnson (1970), 4) Reise (1991), 5) Macnae and Kalk
(1962), 6) Day (1974), 7) Sasekumar (1974), 8) Berry (1964), 9) Vohra (1971), 10) Wells (1983), 11) Piersma et al.
(unpubl.), 12) Pepping et al. (1999), 13) Dittmann (in press), 14) Dittmann (1995), 15) Molina and Vargas (1994),
16) Vargas (1987, unpubl.), 17) Acheampong (2001).

Table 2. Abundances (individuals m2 ) of benthic fauna in tropical tidal flats.


Mean values as well as the minimum and maximum values recorded are
presented. Based on the sieve sizes used, the data are presented for macrofauna
retained on the generally used sieve sizes and on smaller sieve sizes (mesofauna
sensu Dittmann, 1995).
Location

mean

min

max

Macrofauna (0.51 mm sieve)


Banc dArguin, Mauretania 1)
Bay of Rest, West Australia 2)
Roebuck Bay, West Australia 3)
Hinchinbrook Channel, northeast Australia 4)
Haughton estuary, northeast Australia 5)
Punta Morales, Costa Rica 6)

1405
992
1287
1363
1919
14798

152
356
40
57
57
6345

5635
4186
16280
8531
29548
25421

Mesofauna (0.25 mm sieve)


Hinchinbrook Channel, northeast Australia 4)
Haughton estuary, northeast Australia 5)

16080
14140

0
0

59000
59000

1) (Wijnsma et al., 1999), 2) (Wells, 1983), 3) (Pepping et al., 1999), 4)


(Dittmann, in press), 5) (Dittmann, 2000), 6) (Vargas, 1987).

192
Table 3. The five most abundant benthic species (retained on 0.5 or 1 mm sieve) according to a rank order of abundance in various tropical tidal
flats. A = Amphipoda, B = Bivalvia, Br = Brachiopoda, C = Cumacea, D = Decapoda, G = Gastropoda, O = Ophiurida, Os = Ostracoda, P =
Polychaeta.
rank

Banc dArguin,
Mauritania 1)

Roebuck Bay,
Australia 2)

Hinchinbrook
Channel,
Australia 3)

Haughton
estuary,
Australia 4)

Estero de
Jaltepeque,
El Salvador 5)

Punta
Morales,
Costa Rica 6)

1.

Capitella
capitata (P)
Euclymene oerstedii
(P)
Naineris laevigata
(P)

Chaetopteridae
indet. (P)
Oweniidae indet.
(P)
Amphiuridae indet.
(O)

Theodoxus
oualaniensis (G)
Cerithium cf.
coralium (G)
Amygdalum
glaberrima (B)

Dasybranchus
lumbricoides (P)
Cumacea indet.
(C)
Acesta lopezi
(P)

Coricuma
nicoyensis (C)
Mediomastus
californiensis (P)
Paraprionospio
pinnata (P)

Loripes lacteus
(B)
Hydrobia ulvae
(G)

Anodontia cf.
omissa (B)
Glyceridae indet.
(P)

Lingula antina
(Br)
Pitar coxeni (B)

Tellina australis
(B)
Gammaridae
indet. (A)
Armandia
secundariopapillata (P)
Haustoriidae indet.
(A)
Barantolla sp. (P)

Haploscoloplos
elongatus (P)
Sigambra
ocellata (P)

Cyprideis pacifica
(Os)
Pinnixia valerii
(D)

2.
3.

4.
5

1) Wolff et al. (1993), 2) Pepping et al. (1999), 3) Dittmann (in press), 4) Dittmann (2000 and unpubl.), 5) Molina and Vargas (1984), 6) Vargas

(1987).

Spatial distribution patterns


In intertidal environments, hydrodynamic conditions
determine the duration of submergence and distribution of sediment parameters and nutrients, which in
turn determine the spatial distribution of organisms
and even trophic types (Dankers and Beukema 1983,
Peterson 1991). This is well established for temperate locations and distinct spatial zonation in tropical
tidal flats were recorded by Macnae and Kalk (1962),
Berry (1964), Vohra (1971), Day (1974), Sasekumar
(1974), Frith et al. (1976), Gibbs (1978), Morton
and Morton (1983), Warwick and Rushwayuni (1987),
Reise (1991), Schrijvers et al. (1995) and Dittmann
(2000). The occurrence of single species or taxa, feeding types or life forms along the intertidal gradient
is often similar between these studies, with preferences emerging for upper or lower intertidal, muddy
or sandy substrates. Dittmann (2000) analysed the
zonation of benthic communities along an intertidal
gradient, which corresponds with findings from other
Indo-West-Pacific tidal flats.
The distribution of the larger macrobenthic organisms in particular seems to follow a general pattern.
At present this can only be a rough scheme which has
to be substantiated by further quantitative surveys. In
the muddy sediments of the upper intertidal, crabs of
the genus Macrophthalmus occur together with fiddler crabs and other Ocypodidae, mudskippers and
mudwhelks (Cerithiidae, Potamididae). In the mid in-

tertidal, burrowing forms are frequent in muddy sand,


such as Callianassa or other thalassinidean shrimps,
brachiopods (Lingula in the Indo-West Pacific and
Glottidia in Central America), tube worms (of several
polychaete families, e.g. Terebellidae or Onuphidae)
and other surface-deposit-feeders, e.g. echiurids. Bivalves, often of the families Tellinidae or Psammobiidae, can occur in sandflats together with benthic
holothurids. In sandy sediments nearer the lower intertidal, hemichordates are found together with echinoderms like sand dollars or brittle stars. Hermit crabs
can be frequent here, too, or occur throughout the
intertidal.

Ecological roles
Many of the taxa named in the above zonation scheme
modify the substrate or nutrients in such a way that
they become more or less suitable for other species,
and can thus be classified as ecosystem engineers
sensu Jones et al. (1994). In tidal flats, where promotive interactions are known to play an important
role (Reise, 1985), these are common and important
species interactions structuring or regulating benthic
assemblages. In tropical tidal flats, such promotive interactions were described by Macnae and Kalk (1962),
Morton and Morton (1983) and Dittmann (1996).
Burrows of macrobenthic organisms do not only
provide accommodation for smaller infauna, the burrowing activities are often also important for sediment

193
Table 4. Occurrence of several burrow-building benthic taxa in tidal flats at various tropical locations. This list is not complete and gives
only major burrowing organisms. If taxa were not recorded on species or genus level, their occurrence is indicated at a higher taxonomic
level.
Taxa

Crustacea
Uca
Dotilla
Scopimera
Macrophthalmus
Callianassidae
Squilla
Sipunculida
Sipunculus
Siphonosoma
Phascolosoma
Holothuroidea
Echiurida
Brachiopoda
Lingula
Glottidia
Enteropneusta
Saccoglossus
Balanoglossus
Ptychodera
Glossobalanus

Banc

Inhaca

Morrumbene Ao Nam Singapore Hong

Roebuck Anna

dArguin1) Island2) estuary3)

Bor4)

5)

Kong6) Bay7)

Punta

estuary10) Morales11)

Hinchinbrook Haughton

Plains8) Channel9)

1) Mauritania, Wolff et al. (1993), 2) Mozambique, Macnae and Kalk (1962), 3) Mozambique, Day (1974), 4) Thailand, Frith et al.
(1976), Reise (1991), 5) Vohra (1971), 6) Morton and Morton (1983), 7) Pepping et al. (1999) 8) Piersma et al. (1999), 9) Dittmann
(1995, in press and unpubl.), 10) Dittmann (1995, 2000 and unpubl.), 11) Vargas (1987 and unpubl.).

destabilisation, bioturbation and benthic-pelagic coupling. Taking burrowers as an example for an ecological role, the occurrence of burrowing taxa in tropical
tidal flats has been compiled in Table 4. Because of
taxonomic uncertainties, this comparison is not carried
out on a species level. As Table 4 shows, similar taxa
occur at various sites and certain genera are especially
widely distributed in the Indo-West Pacific. Each site
has a site-specific mix of burrowers. At some sites like
Inhaca Island, a great many burrow-building species
co-exist, including closely-related taxa.
The presence or absence of species or taxa
with certain ecological functions can result in sitespecific variations in processes structuring or regulating benthic assemblages. I hypothesize that a greater
habitat diversity in tropical tidal flats determines sitespecific occurrences of ecosystem engineers, which
accounts for the greater variation in benthic assemblages between tropical tidal flats (Dittmann and
Vargas, 2001). A comparison of taxa responsible for

certain ecological roles in tidal flats of northern Australia and Central America revealed that mainly related taxa fill similar ecological niches (Dittmann and
Vargas, 2001).

Conclusions
This literature survey was set out to assess possible
variations in benthic assemblages of tropical tidal flats
to facilitate future global comparisons. The analysis
shows that most tropical tidal flats are rich in species
and low in individual abundances. Species occurrence
appears to be site-specific, in spite of similarities in
spatial distribution patterns, especially in the IndoWest Pacific. Site-specific idiosyncrasies in the presence or absence of certain ecosystem engineers can
account for variations in benthic assemblages between
sites. Therefore, processes structuring and regulating benthic assemblages can vary between tropical
tidal flats. Such singularities have applied implica-

194
tions, as they complicate generalizations for impact
assessments. Using the same methodology, further
biogeographic comparisons of intertidal assemblages
should be carried out in the future.

Acknowledgements
I thank the organizers of the MMM conference for an
inspiring meeting and the opportunity to see tidal flats
on the coast of Kenya.

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