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Journal of Fish Biology (2002) 61, (Supplement A), 1–20

doi:10.1006/jfbi.2002.2063, available online at http://www.idealibrary.com on
‘ Fish in hot water ’: the challenges facing fish and fisheries
research in tropical estuaries
S. J. M. Bi\nr×
CSIRO Marine Research, P.O. Box 120, Cleveland, Queensland 4163, Australia
Tropical estuarine areas comprise small systems of a few km, larger estuaries, coastal lakes of
hundreds of km
2
and vast shallow coastal waters that are contiguous with estuaries and have
similar reduced salinities. Many of the world’s great estuaries are in the tropics, e.g. the
Amazon, Orinoco, Congo, Zambezi, Niger, Ganges and Mekong. The distribution of tropical
and subtropical estuaries approximately follows that of mangroves. Like estuaries everywhere,
they are a focus of human activity and are among the most exploited of ecosystems. In few
other places do the activities of fishers, industrialists, shippers, farmers, conservationists, sports
enthusiasts and biologists overlap to such an extent. Quite apart from the possible effects of all
these activities, the fishes of subtropical and tropical estuaries already face one of the most
rigorous of aquatic environments; but despite this, species diversity and productivity are high.
Only in tropical estuaries are animals from such a wide range of taxa so closely associated,
annelid worms, prawns, crocodiles, birds, ‘ hippos ’, dolphins and of course fishes, all may form
part of the overall community, often with functional ecological links. Unfortunately, the
difficulties of working in these often inhospitable environments, has meant that biologists have
favoured projects in more appealing areas, such as coral reefs. While it is still true that most
estuarine research is conducted in industrialized developed countries, nearly all of which are in
cold or temperate regions, there has been a recent upsurge in tropical estuarine fish research.
This is being driven by two imperatives, food security and the conservation and maintenance of
biodiversity. Both these problems require knowledge of the ecology of tropical estuarine fishes,
particularly their relationships with the environment and the extent to which they are dependent
on estuaries or adjacent habitats for survival.
2002 The Fisheries Society of the British Isles. Published by Elsevier Science Ltd. All rights reserved.
Key words: fisheries; tropical estuaries; research; fish; biodiversity.
INTRODUCTION
This review considers fishes living in tropical and sub-tropical estuaries where the
water temperature seldom falls below c. 25 C, but these environments are also
the focus of a great variety of human activities, many inimical to the survival of
fishes! They already face one of the most rigorous of aquatic environments, with
many species operating at or close to their physiological limits and exhibiting a
large array of ecological specializations. Unfortunately, their ultimate fate may
not be determined only by these adaptations, but by the ways in which humans
use and mis-use estuaries.
To set the scene, it is necessary to summarize: the variety of estuaries in the
tropics and their myriad habitats; to briefly describe the great diversity of their
fishes; the similarities between the fish faunas of estuaries often far apart; the
physical and biological factors that help determine their diversity. Against a
background of rapidly expanding knowledge of tropical estuarine fishes, it is
necessary to consider what are the research challenges, biological, physical and
logistic, which have to be overcome if, for example, some of the contentious
Tel.: +61 7 38267214; fax: +61 7 38267281; email: steve.blaber@csiro.au
1
0022–1112/02/61A001+20 $35.00/0 2002 The Fisheries Society of the British Isles. Published by Elsevier Science Ltd. All rights reserved.
paradigms such as those related to estuarine dependence or the importance of
mangroves to fisheries production have to be dealt with. In addition, no
discussion of tropical estuarine fishes can ignore the human dimension, both in
terms of fishing and the range of other activities that may impact on the fishes.
THE ESTUARIES
Many of the world’s great estuaries are in the tropics (the Amazon, Orinoco,
Congo, Zambezi, Niger, Ganges and Mekong) and are all very large and receive
drainage from enormous catchments. Tropical estuarine environments, how-
ever, also include tiny seasonally flowing systems of 1 to 2 km. They also
encompass extensive coastal lakes and the reduced salinity estuarine waters
extending along the coast in parts of South East Asia, South America and
Africa. There is thus a lack of uniformity among tropical estuaries in terms of
size, depth, habitats and physical regimes. These factors and many others, such
as the nature of the adjacent marine and freshwater habitats, can greatly
influence the fish faunas and fisheries of tropical estuaries.
The world-wide extent of tropical and subtropical estuaries approximately
follows the distribution of mangroves, which are the dominant intertidal
vegetation in subtropical and tropical estuaries (Chapman, 1976; Duke, 1992).
Mangroves generally match the 20 C isotherms in both hemispheres, suggesting
that water temperature is the most significant influence. Hutchings & Saenger
(1987) concluded that the presence of mangroves correlates with areas where the
water temperature of the warmest month is >24 C; also that their northern and
southern limits correlate reasonably well with the 16 C isotherm for the air
temperature of the coldest month. Latitudinal ranges are greater on eastern
continental margins than on western sides due to the presence of warm or cold
currents. Tropical estuaries grade into subtropical systems beyond the tropics of
Cancer and Capricorn where a winter water temperature low of c. 12 C marks
their southern and northern limits (Whitfield, 1994; Ayvazian & Hyndes, 1995).
Although many estuaries of the south and south east U.S.A. have been described
as ‘ tropical ’ or ‘ subtropical ’ (McPherson & Miller, 1987; Powell et al., 1991),
this is not really the case in a world context, with the possible exception of south
Florida mangrove-lined systems. Winter water temperatures in many U.S. Gulf
of Mexico estuaries fall as low as 5 C (Deegan & Thompson, 1985).
To facilitate the description of those factors most relevant to their fish and
fisheries, tropical estuaries can be divided into four broad categories: open
estuary, estuarine coastal waters, coastal lake and blind estuary (the latter are
sometimes referred to as lagoons, but this term is best avoided as it is also used
in a number of other contexts, e.g. in relation to coral reefs). It must however be
emphasized that any divisions are for convenience and a continuum exists.
OPEN ESTUARIES
This category includes all the larger well-known estuaries of most of the
medium and larger rivers of the tropics and comes closest to the classical
European and North American concept of an estuary (Elliott & McLusky, in
press). These estuaries are never isolated from the sea and are subject to tidal
influence with all the physical consequences of regular salinity, turbidity and
2 s. ¡. x. ni\nr×
current flow changes, tidal prisms, haloclines and intertidal habitats. Extensive
deltas occur at the mouths of many of the larger estuaries, such as those of the
Zambezi and Niger in Africa, the Orinoco in South America and the Mekong in
Asia, and the largest of them all, the Ganges (Meghna) in India and Bangladesh.
The size and depth of this sort of estuary has made them a focus of human
activity, including ports and harbours with their associated industrial and
city developments and consequent effluent problems. Fishery activities have
increased in many of these estuaries, often developing from a subsistence base,
through an artisanal phase to fully-fledged commercial operations, to supply the
demand from burgeoning cities. This increases the pressure on fish resources and
the effects of fishing may lead to changes in the fish fauna and fish community
structure.
ESTUARINE COASTAL WATERS
Nowhere is the problem of definitions and boundaries of estuaries and
estuarine fish faunas better illustrated than in tropical coastal waters. Many of
the larger open estuaries grade almost imperceptibly into the adjacent sea. The
effects of the discharge from the Amazon are felt up to 400 km from the mouth
and those of the Orinoco up to 50 km away from the mouth. The shallow nature
of such tropical coastal waters and their lowered salinities and high turbidities
make them at least partly estuarine in character, particularly as regards their fish
faunas. The cut-off point between these waters and true marine conditions
may best be defined by the fauna (Pauly, 1985) rather than by physical factors,
but estuarine conditions (defined by salinity) seldom extend beyond depths of
c. 15–20 m, often related to the thermocline depth. For example, off Guyana
these waters extend 40 km from shore (Lowe-McConnell, 1987), in the Gulf of
Carpentaria c. 10 km from shore (Blaber et al., 1990), in the South China sea
20–50 km from the Borneo coast and in the Bay of Bengal >100 km from the
mouth of the Ganges. Most estuarine coastal waters are subject to heavy fishing
pressure, mainly from trawlers.
COASTAL LAKES
Many lacustrine bodies of water behind tropical shorelines on most continents
are estuarine in character. Their faunas are mainly marine or estuarine, and
most coastal lakes have some form of connection to the sea. The form and
constancy of this connection largely determines the prevailing salinities and the
nature of the fish fauna. The range of habitats is also determined by this
connection as well as by the bathymetry, which is a result of local geological
history and processes. They are sometimes referred to as coastal lagoons
(Barnes, 1980), but the term lake is used to avoid confusion with small lagoons
found at the mouths of blind estuaries. What makes these systems unique is their
relatively large surface area. Many have been extensively modified by man and
most have important fisheries, recreational and conservation value. Important
chains of coastal lakes occur along the south-east and west African coasts, in
India, in the Gulf of Mexico and in northern South America.
BLIND ESTUARIES
The most useful definition of blind or temporarily closed estuaries is that of
Day (1981) who described them in the following terms: ‘ These are estuaries that
risn iN no1 v\1r× 3
are temporarily closed by a sand bar across the sea mouth. At such times there
is no tidal range and thus no tidal currents. Freshwater enters from the river and
the circulation is dependent on the residual river current and the stress of the
wind on the water surface. According to the ratio between evaporation and
seepage through the bar on the one hand, and freshwater inflow plus precipita-
tion on the other, the salinity will vary. ’ Estuaries in this category are usually
relatively small, both in length and catchment and are characteristic of areas of
low or highly seasonal rainfall. Although best developed along the warm
temperate and Mediterranean climate coasts of southern Australia and South
Africa (Potter et al., 1990), they do extend into the sub-tropics and tropics of
both Africa and India. In addition, equatorial examples occur in South East
Asia. Because of their relatively small size, blind estuaries are not often
subject to commercial fishing pressure, but are usually intensively exploited by
subsistence fishers.
THE FISHES
The species richness of fishes in tropical estuaries is greatest in the Indo-West
Pacific, but larger systems usually have more species than smaller ones; deep
open water channels in the larger systems favour more of the larger species,
particularly carangids and sharks, in addition to a higher number of occasional
visitors. In addition, larger systems usually have a greater diversity of habitats.
Within the tropics, it is therefore a combination of estuary size and diversity of
habitats that are the most important factors in relation to species diversity.
Hence comparisons of species richness between estuaries of the Indo-West
Pacific and the East and West Tropical Atlantic should be viewed with caution.
Taking into account the large size of some Atlantic systems (e.g. Orinoco
estuary, Termino´ s and Lagos Lagoons), however, it is apparent that even small
estuaries in the Indo-West Pacific are usually more species rich than Atlantic
systems (Table I). Comparisons of overall fish biomasses in different tropical
estuaries are difficult because there are few published figures, and also because
many of the results may not be comparable due to differences in methods. The
scarce data (Table II) suggest, however that fish biomass is not usually >30 g
m
2
and is most commonly between c. 5 and 15 g m
2
. These values are an
order of magnitude higher than those of tropical non-estuarine seagrass areas in
shallow waters (Blaber et al., 1992).
The number of fish species in subtropical and tropical estuaries is much greater
than in temperate regions. Where individual temperate estuaries may have c. 20
commonly taken species [ignoring rare and adventitious species (Potter et al.,
1986; Elliott & Taylor, 1989; Pomfret et al., 1991; Elliott & Hemingway, 2002)]
and warm temperate ones c. 50 (Darnell, 1961; Lenanton & Hodgkin, 1985),
most medium to large subtropical and tropical estuarine areas have at least 100,
with some reaching >200. Additionally, the number of species in adjacent
estuarine coastal areas often exceeds 300, but depends upon the presence of
specific habitats such as seagrass beds. The composition of the fish faunas results
from the interplay of a whole range of factors among which the most important
are: (a) Estuary size, depth and physical regimes, particularly salinity and
turbidity, as well as the types of habitats, particularly the composition and extent
4 s. ¡. x. ni\nr×
of mangroves; (b) the nature and depth of adjacent marine waters and to a much
lesser extent, fresh waters; (c) the geographical location of the estuary, both in
terms of latitude and in relation to marine features such as ocean currents,
canyons and reefs. It is also important to emphasize that the number of species
recorded in any system is a result of the extent of sampling, both spatial and
temporal.
T\nir I. Numbers of species recorded from different types of estuaries (O, open; B, blind;
CL, coastal lake) in the four major zoogeographic regions (modified from Blaber, 2000)
System Country Type and size
Number of
species
Indo-west Pacific
Alligator Creek Australia O small 150
Trinity Australia O medium 91
Embley Australia O large 197
Vellar Coleroon India O large 195
Chilka India CL large 152
Hooghly India O large 172
Chuwei Taiwan O small 30
Ponggol Singapore O medium 78
Matang W. Malaysia O medium 117
Kretam Kechil E. Malaysia O small 44
Purari New Guinea O large 140
Pagbilao Philippines O medium 128
Solomon Islands Solomon Islands* O small 136*
Morrumbene Mocambique O medium 113
Tudor Creek Kenya O small 83
Gazi Bay Kenya CL small 128
Kosi South Africa CL large 163
St Lucia South Africa CL large 110
Mhlanga South Africa B Small 47
East Atlantic
Lagos Nigeria CL large 79
Fatala Guinea O medium 102
Ebrie´ Ivory Coast CL large 153
Niger Nigeria O large 52
West Atlantic
Guaratuba Brazil CL medium 61
Itamaraca Brazil O large 81
Orinoco Venezuela O large 87
Terminos Mexico CL large 122
Cienaga Grande Colombia CL large 114
Tortuguero Costa Rica O small 70
Sinnamary French Guiana O medium 83
East Pacific
Rio Claro Costa Rica O small 22
Guerrero Lakes Mexico** CL small 105
*Incorporates 13 small estuaries (no one system more than 50 species).
**Sum of species for 10 small coastal lakes.
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The classification of estuarine fishes has been almost as numerous as the
classification of estuaries (Day, 1951; Gunter, 1967; McHugh, 1967; Green,
1968; Perkins, 1974; Whitfield, 1998) with taxonomic, physiological and
ecological groupings based on attributes such as salinity tolerance, breeding,
feeding and migratory habits. Most research on estuarine fishes has been in
temperate and warm temperate regions of Europe and North America where
salinity has long been regarded as a key factor regulating the composition of
estuarine fish faunas. Gunter (1967) considered the lower salinity of estuarine
waters to be the outstanding difference between these waters and sea water.
While this phenomenon has been amply demonstrated for warm temperate and
temperate areas it is not at all clear in the subtropics and tropics. Salinity is only
one of an array of factors that determine which species are found in any one
subtropical or tropical estuary. Salinity tolerance may play a role in the
distribution of fishes in tropical estuaries, but as these systems often undergo
very great fluctuations in salinity (often from almost 0 to 35), both diurnally and
seasonally, a considerable degree of euryhalinity is a pre-condition and pre-
requisite for their inhabitants, as well as for many species in marine areas that
suffer seasonal reductions or fluctuations in salinity.
For this reason, a grouping of tropical estuarine fishes based mainly on their
salinity tolerances is not very useful or practical, especially as there has been little
experimental work conducted on the salinity tolerances of most species. Based
on their occurrence in low salinity waters almost all fit into a similar, very
euryhaline category. This has led to a number of classification systems for
subtropical and tropical estuarine fishes (Day, 1981; Blaber, 1997; Whitfield,
1998), based mainly on how they utilize estuaries and where they spawn. The
following classification scheme is similar, but gives increased importance to the
‘ estuarine ’ category in the tropics due to recent advances in knowledge of
spawning behaviour (Blaber, 2000):
(a) Estuarine species that can complete their entire life cycle in the estuary.
Sub-tropics: they represent a small but significant part of the estuarine fish
fauna and include a number of clupeids (e.g. Gilchristella aestuaria
(Gilchrist) in Africa), gobies (hundreds of species), engraulids (e.g. several
species of Coilia in South East Asia), ambassids, atherinids and
syngnathids. Tropics: they represent a large part of the estuarine fish
fauna, particularly in West Africa, South East Asia and tropical South
America, and include many species of Clupeidae, Mugilidae, Sciaenidae,
Ariidae, Polynemidae and Haemulidae that spawn in estuaries.
(b) Marine migrants from the sea, usually characterized as marine spawners.
This is a large group in subtropical and tropical estuaries where they
may occur both as juveniles and adults [e.g. Lates calcarifer (Bloch) of
South East Asia and Australia; the circum-tropical Mugil cephalus L.,
as well as most Leiognathidae and Haemulidae], or only as juveniles
(e.g. some species of Mugilidae, Carangidae and Polynemidae) or only
as adults (e.g. some species of Ariidae). It is important to note,
however, that the division of this category from that of (a) is often
difficult in the tropics, because many of the species that fall into the
marine migrant category in the sub-tropics, form part of the ‘ estuarine ’
category in the tropics.
risn iN no1 v\1r× 7
(c) Anadromous species that breed in fresh water and on their way to and
from their spawning grounds spend time in estuaries. This group is
important in temperate estuaries, but is a rare category in subtropical and
tropical estuaries, but includes the commercially important tropical shads
Tenualosa ilisha (Hamilton) and Tenualosa toli (Valenciennes) of south
Asia and Sarawak respectively. Similarly, catadromous species are rare in
the tropics
(d) Freshwater migrants that move varying distances down estuaries, some-
times to spawn (e.g. eleotrids), but usually return to fresh water to breed.
In the tropics this group is best represented in South and Central
American estuaries where a number of pimelodid catfish, poeciliids and
characids penetrate estuarine waters. The freshwater cichlid Oreochromis
mossambicus Peters and the archerfishes (Toxotidae) are well known
African and South East Asian examples.
The fish communities of subtropical and tropical estuaries of all four major
tropical zoogeographic regions (Indo-West Pacific, East Atlantic, West Atlantic
and East Pacific) have many common characteristics. In almost all cases, they
are dominated by fishes of marine origin, with more than half the number of
species as well as the number of individuals, being contributed by either fully
estuarine species or marine migrants. Given the ‘ estuarization ’ of many tropical
coastal areas, this is not surprising and perhaps the term ‘ marine ’ for many of
these species is a misnomer, for many of them do not occur outside of estuarine
coastal areas. The dominant taxa in each region are broadly similar, but there
are some interesting contrasts. In all regions except the Indo-West Pacific,
Sciaenidae are one of the dominant families. In the Indo-West Pacific, sciaenids
are important in the equatorial regions of South East Asia, but much less so
elsewhere. This pattern may be connected with the amount of rainfall and the
degree of estuarization of coastal waters. In East Africa for example, there are
few estuarine coastal waters and sciaenids, and although present they are not a
dominant component of the fauna. Except for the sciaenids, the other dominant
families are broadly comparable across all regions. It is noteworthy, however,
that within the Indo-West Pacific, clupeids and engraulids are much more diverse
and numerous in the estuarine coastal waters of equatorial South East Asia than
in other areas.
THE CHALLENGES
THE STATUS OF TROPICAL ESTUARINE FISH RESEARCH
While it is still true that most estuarine research is conducted in industrialized
developed countries, nearly all of which are in cold or temperate regions, there
has been an upsurge in tropical estuarine fish research in the last 5 years. This is
being driven by two imperatives: firstly that of the plight of inshore fisheries in
developing countries, where declining catches, environmental degradation and
increasing human populations are jeopardizing food security, and secondly
under pressure, largely from the developed world, for conservation and main-
tenance of biodiversity. Both these problems require detailed knowledge of the
ecology of tropical estuarine fishes, particularly their relationships with the
environment and the extent to which they are dependent on estuaries or adjacent
habitats for survival. A search of the Aquatic Science and Fisheries Abstracts

8 s. ¡. x. ni\nr×
database from 1978 to the end of 1995 revealed c. 1800 publications dealing with
fishes in temperate estuaries, whereas there were only 600 for tropical estuarine
fishes. For the period from 1995 until the present the figures are 630 for
temperate estuarine fishes and 338 for tropical estuarine fishes, a considerable
increase in productivity from the tropics.
BIOLOGICAL
Recent advances in the knowledge of tropical estuarine fish ecology cover most
aspects of the science. A great deal more basic data on species composition and
diversity of systems are now available. This is particularly the case in South and
West Africa, where syntheses of much of the previous research are being
produced (Whitfield, 1998; Albaret, 1999); South America, where the results of
recent research in Brazil (Chaves & Bouchereau, 1999) and Colombia (Rueda &
Santos-Martinez, 1999) are appearing; throughout South East Asia, where
increasing affluence and levels of education have added to many both national
and international research efforts. Detailed data on the feeding and reproduc-
tion of many species are becoming available, much of them very relevant to
understanding ecological relationships in relation to various forms of exploita-
tion and environmental changes. For example, the discovery that several of the
tropical clupeid shads (genus Tenualosa) are protandrous hermaphrodites has
profound implications for both their fisheries management and conservation
(Blaber et al., in press). The role of the mangrove environment, primarily as a
nursery for juvenile fishes and prawns, has been debated for a long time.
Following some major research efforts, the complex functional relationships are
now becoming evident (Chong & Choo, 1999) and ways of quantifying the
relationship between mangroves and fisheries production are being seriously
examined and studied (Baran, 1999).
Unfortunately, one of the major obstacles to thorough ecological research still
remains the difficulties associated with the correct taxonomic identification of
tropical marine and estuarine fishes. Although a number of very useful new field
guides are now available (Allen, 1997), and new FAO guides are appearing, the
large number of species and lack of adequate field keys frequently hamper
research, particularly in developing countries where scientists seldom have access
to good library facilities. A more worrying aspect is the overall decline in
funding throughout the world for taxonomic research, the decline in university
teaching of taxonomy, and consequently the very real decline in numbers of fish
taxonomists. Other important practical biological problems, particularly in
developing countries, include appropriate sampling design, accurate sampling
and statistical analysis.
PHYSICAL
One of the most difficult challenges is to separate and evaluate the influences
of each a suite of physical parameters influencing estuarine fishes. Many are
correlated with one another. In terms of hydrology, current speed and turbidity,
and to a lesser extent, salinity, all are important. Turbidity has been shown to be
an important determinant of species composition in estuaries (Blaber & Blaber,
1980; Cyrus & Blaber, 1987), but the effects of current speed are not well
understood. Current flow in tropical estuaries, such as the Norman in northern
risn iN no1 v\1r× 9
Australia (Blaber et al., 1994), results largely from tidal range and freshwater
inflow. Both are also correlated with highly variable salinities and high
turbidities. Further evidence of the relative importance and interrelationships of
physical factors comes from work in the Sarodrano mangroves, in a semi-arid
area of Madagascar (Laroche et al., 1997). Here, temperature and salinity have
no effect on fish community structure, and the turbidity gradient is constant
throughout the year and does not cause any seasonal variations in the distribu-
tion of juveniles. Most of the variation in species composition is correlated with
tidal range, lunar phase and time of day or night.
In addition, habitat diversity and structure are important in relation to species
composition; within the Gulf of Carpentaria in northern Australia, comparing
sites of similar size, the Embley Estuary, with a broad range of habitats,
including a wide variety of mangroves, has more species (197) than either Groote
Eylandt (179) or the relatively homogeneous Norman Estuary (100) which has a
narrow band of fringing mangroves. The influence of the structural heterogen-
eity of each habitat is important, particularly in mangroves (Thayer et al., 1987);
this is reflected by the greater number of species found in the complex forests of
the Embley than in the more depauperate Norman Estuary. It is thus probable
that the fish communities of estuarine and inshore waters of the tropical
Indo-West Pacific are at least partly determined by the interrelationships
between current speed, tidal range, turbidity and salinity, as well as by the
structure of the habitat. The relative importance of each factor varies
from estuary to estuary, but the variable combinations produce communities
characteristic of particular conditions.
LOGISTIC
Tropical estuaries contain a wide range of taxa that are very closely associated,
annelid worms, prawns, crocodiles, birds, ‘ hippos ’ and of course fishes, all may
form part of the overall tropical estuarine community, often with functional
ecological links. Unfortunately, the difficulties of working in these often
inhospitable, usually muddy and hot environments with many biting insects, has
meant that biologists have tended to favour projects in more appealing areas,
such as coral reefs.
Much of the research on fishes in tropical estuaries is done in developing
countries, usually for important fisheries purposes, often linked to poverty
alleviation, and not necessarily for fundamental scientific reasons. Resources
and facilities in such areas are usually limited or sometimes non-existent and
publication of results, whether or not they are important, may not occur. Many
of the results are only to be found in the ‘ grey literature ’ of annual reports,
reports to aid or funding organizations, unpublished conference papers and
articles in newsletters. As stated by Blaber (2000), such data are often important
and may be the only information on a particular area or species and it is vital to
draw fully on this literature to support and extend results from the primary
literature.
POPULAR PARADIGMS—ESTUARINE DEPENDENCE
Whether or not particular species, populations, communities or certain life
history phases of fishes that occur in estuaries are dependent on the estuarine
10 s. ¡. x. ni\nr×
environment for feeding, refugia or spawning, as opposed to the sea or even fresh
water, has given rise to much discussion and speculation (Hedgpeth, 1982).
Unfortunately there is little agreement in general terms about the validity of
‘ estuarine dependence ’, due mainly to parochialism, and the all too ready
acceptance by workers everywhere of the paradigm developed for fishes of the
estuaries of the south eastern U.S.A. (Gunter, 1967; McHugh, 1967; Day &
Ya´n˜ ez-Arancibia, 1985). The concept of an estuarine dependent phase in the life
cycle of coastal fishes arose primarily from studies of temperate and warm
temperate estuaries in the U.S.A. and in southern Africa. In these non-tropical
areas many marine species have been shown to be dependent on the estuarine
environment during their juvenile phase (Day et al., 1981; Deegan & Thompson,
1985). For example in the estuaries of the Louisiana deltaic plain, species
utilizing the estuary as a nursery make up 35 to 72% of all species, and usually
over 90% of individuals (Deegan & Thompson, 1985).
The degree of estuarine dependence among fishes of the sub-tropics and
tropics was questioned by Longhurst & Pauly (1987) who analysed studies from
20 subtropical and tropical areas. They concluded that only in Brazil and South
Africa, in systems that are marginally subtropical, could estuarine dependence
really be demonstrated. Blaber (1981) and Blaber & Blaber (1980) postulated
that most of the estuarine fish fauna of the sub-tropics and tropics is not
‘ estuarine ’ per se, but is a fauna characteristic of shallow, turbid areas, often
with variable salinity. Areas with such characteristics occur over very large areas
of the sea in South and South East Asia, West Africa and northern South
America, but are confined mainly to estuaries in southern Africa, and parts of
the U.S.A. and Australia. Many of the taxa considered largely estuarine
dependent in, for example, South Africa, such as Gerres, Thryssa and Elops are
common in estuarine coastal waters throughout South and South East Asia.
In southern Africa, Whitfield (1994) divided the 142 estuarine fishes into five
categories in relation to estuarine dependence: estuarine species (species that
breed in estuaries, 28%), euryhaline marine species (species that breed in the sea
and juveniles occur in estuaries, 43%), marine species (occur in estuaries in small
numbers as occasional visitors, 21%), euryhaline freshwater species (may breed
in estuaries, 5%), and obligate catadromous species (species that use estuaries in
transit between the sea and fresh water, 3%). Therefore according to this
classification 71% of the species are completely or partially dependent on
estuaries. It is important to note, however, that this classification includes a
mixture of temperate and warm temperate species as well as those of subtropical
South East Africa; many of the subtropical species included in this classification
are at the southern limits of their distribution, and as discussed earlier, are not
strictly estuarine dependent in the tropics.
The region with the greatest diversity of coastal fish species is the tropical
Indo-West Pacific, where vast areas of estuarine coastal waters in South and
South East Asia, south to parts of northern Australia, form what Blaber (1997)
termed the ‘ core area ’ for the majority of Indo-West Pacific estuarine species.
Briggs (1974) proposed that the tropical Indo-Malayan region was the centre of
speciation among Indo-West Pacific shallow water fishes, with its unparalleled
variety of habitats, from coral reefs to seagrasses to mangroves and estuaries.
Whether the greater number of species in this region is due to enhanced
risn iN no1 v\1r× 11
speciation rates or reduced extinction rates remains a subject of controversy
(Briggs, 1974). Nevertheless, the similarity of tropical estuarine fish faunas
across the great breadth of the Indo-West Pacific is striking. For example, at a
species level, two thirds of the species in the St Lucia coastal lake system of
KwaZulu-Natal, South Africa and 71% of species in Madagascar estuaries occur
in South East Asia. Almost all species in the estuaries of northern Australia are
also common throughout South and South East Asia. For most families there is
also a gradual reduction in species diversity away from the ‘ core area ’,
particularly where physical conditions change and mangrove habitats are not
dominant (Blaber, 2000). Such marginal areas are restricted to the relatively
small areas of estuaries along south-east African, east Australian and south
Pacific shores where shallow, turbid and variable salinity conditions give way to
coral reefs and clearer waters.
The majority of species found in tropical estuaries are probably not estuarine
dependent, but the question of total estuarine dependence may only be relevant
in estuaries outside the ‘ core areas ’. It has less relevance where there is
extensive estuarization (Longhurst & Pauly, 1987) of the continental shelf, where
most species remain in similar (large and widespread) habitats for their whole
life. Many more species spawn in tropical estuarine areas, including particularly
the Sciaenidae, Engraulididae and Clupeidae, than they do in temperate or warm
temperate estuaries.
Resolving the question of the importance of ‘ estuarine dependence ’ for
tropical fishes is hampered because the interrelationships between environmental
factors, habitats, and fish community structure have not been worked out
fully. Here perhaps, lies the main research challenge in relation to ‘ estuarine
dependence ’ in tropical fishes.
POPULAR PARADIGMS—MANGROVES AND FISHERIES PRODUCTION
The degree to which the size and productivity of coastal and offshore fisheries
in the tropics is dependent on the extent of healthy nursery or feeding areas in
estuaries or mangroves has given rise to much debate. Interesting comparative
information is available for some species of penaeid prawns that like many fish
species, have a juvenile phase that lives in estuaries and mangroves. Research in
the Gulf of Mexico (Turner, 1977), Indonesia (Martosubroto & Naamin, 1977),
Australia (Staples et al., 1985) and the Philippines (Primavera, 1998), provides
good evidence that there is a correlation between commercial offshore prawn
catches and the total area of adjacent mangroves. Pauly & Ingles (1986)
concluded that most of the variance in the catches of penaeids could be explained
by a combination of mangrove area and latitude.
The situation with regard to fishes is less clear, and led to Robertson & Blaber
(1992) concluding, that in spite of apparent evidence of a correlation between
mangrove area and commercial fish catches, a causal link has not been
established experimentally. Certainly, Yanez-Arancibia et al. (1985) showed a
positive correlation between commercial fish catches and mangrove area in the
Gulf of Mexico, and Barbier & Strand (1998) determined the effects of changes
in mangrove area in the Laguna de Terminos (Gulf of Mexico) on the production
and value of prawn harvests in Campeche from 1980 to 1990. In southern
Vietnam, De Graaf & Xuan (1998) demonstrated a similar relationship. In the
12 s. ¡. x. ni\nr×
latter study, it was calculated that one hectare of mangrove forest supports a
marine catch of 450 kg year
1
. The few studies that have quantified relation-
ships between mangroves and coastal resources (mainly penaeid prawns) were
summarized by Baran (1999) (Table III), but it must again be reiterated that
these are correlations and that causal links have not been established experimen-
tally. Unfortunately, as stated by Baran (1999) and Baran & Hambrey (1999), in
recent important reviews of the subject, all the studies to date suffer from
problems of auto-correlation, with many factors other than just mangrove area,
such as river discharge, area of shallow coastal water, intertidal area and food
availability, contributing to the relationships.
There is also no general agreement about whether any such relationship
between mangrove area and fisheries production is linear. It has been suggested
that much of the functional value of mangroves might be retained from a smaller
area of mangroves, for example, 75% of nursery function from 50% of original
area (Kapetsky, 1985). It is possible that the mangrove forests fringing deeper
water contain much of the functionality compared with the more inland
shallower, intertidal areas (Vance et al., 1996). Hence, if much of the loss of
mangroves could be confined to the inland side, and deeper fringing areas left
intact, perhaps much of the functional value could be retained. Unfortunately,
it is these deeper fringing areas of mangroves adjacent to the sea that have been
most attractive and suitable for aquaculture pond development throughout the
tropics.
The calculation of any historical relationships between fisheries production
and mangrove area are complicated by changes in both fishing effort and fishing
methods. Overall declines in fish catches in many parts of the tropics coincide,
not only with massive reductions of mangrove area, but also with great increases
in fishing effort, and improved fishing technology. Also, where any increases in
fishing effort are concentrated on juveniles this may affect adult (fishable) stocks.
Hence the ability to use any time-series data to quantify fisheries-mangrove
relationships is compromised.
Baran & Hambrey (1999) discussed the economic value of the fisheries
function of mangroves and indicate that most published monetary values are
misleading because they use aggregated fisheries production and uniformity of
mangrove nursery function. In fact, as stated by Robertson & Blaber (1992) and
Baran & Hambrey (1999), the nursery function of mangroves is highly variable
at local, regional and global scales. Only by using accurate production figures
and a suite of ecological data for each species assemblage, would it be possible to
provide meaningful economic values (Baran & Hambrey, 1999). These authors
conclude that the integration of economic, physical and biological data offers the
only hope for arriving at both an understanding of the fisheries and mangrove
relationship, and finally at meaningful economic evaluations.
THE HUMAN DIMENSION
From a fisheries research or management perspective, taking account of the
activity of people in estuaries is perhaps the hardest challenge of all, especially
for biologists. It requires the integration of a number of disciplines and inputs,
including scientific, sociological (including cultural) and economic. Tropical
estuaries are among the most exploited ecosystems in the world. As the focal
risn iN no1 v\1r× 13
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point for much human activity at the meeting place of sea and river, they have
provided quiet and sheltered waters for harbours, and historically gave the
easiest or only access to the interior for trade or settlement. For peoples living
in the interior they have acted as a route to the sea for trade, fishing and
migration.
Although they may have the highest economic value per hectare of any aquatic
environment (Costanza, 1997), they also have some of the most intractable
problems. Most large tropical or subtropical estuaries have port cities associated
with them that usually have major industries that pour their effluents into the
estuary, which together with discharges from shipping, have resulted in the lower
reaches of many estuaries becoming badly polluted. Industries bring population
increases and growth of cities, which discharge their sewage into the estuary.
Nevertheless, most tropical estuaries are still vitally important as a source of
fishes to the local people and fishing plays an important role in the economy
(Table IV). Hence the overriding need to reconcile conflicts between uses and
users. The effects of fishing itself is a subject that is receiving much attention
throughout the world (Hall, 1999; Blaber et al., 2000). Such effects in estuaries
include not just overfishing, but also alterations to, or removal of the habitat and
the indirect effects of changing fish community structure by selectively removing
some species. Allied to the fishing industry is that of aquaculture. The
development of large-scale aquaculture ventures in tropical estuaries has usually
been at the expense of mangroves and ultimately wild fisheries production.
T\nir IV. Production of fish in different types of tropical estuaries (modified from
Blaber, 2000)
Country Estuary Type T km
2
India Hooghly-Matlah Open estuary system 11·4
Vellar-Coloroon Open estuary system 11·1
Malaysia Larut-Matang Open estuary system 38·64*
U.S.A. Texas bays Estuarine coastal waters 12·1
El Salvador Jiquilisco Estuarine coastal waters 1·7
Philippines San Miguel Bay Estuarine coastal waters 23·8**
South Africa Kosi system Coastal lakes and open estuary 1·0
Ivory Coast Eubrie´ Lagoon Coastal lake 16·0
India Lake Chilka Coastal lake 3·7
Lake Pulicat Coastal lake 2·6
Mandapam Lagoon Coastal lake 5·6
Ghana Sakumo Lagoon Coastal lake 15·0
Malagasy Pangalanes Lagoon Coastal lake 3·7
Colombia Cienaga Grande Coastal lake 12·0
Mexico Caimanero Lagoon Coastal lake 34·5
Terminos lagoon Coastal lake 20·0
Tamiahua Lagoon Coastal lake 4·7
Venezuela Lake Maracaibo Coastal lake 1·9
Tacarigua Lagoon Coastal lake 11·0
*Includes penaeid prawn catches and not coastal waters.
**Probably an overestimate as trawlable biomass only 2·13 t km
2
.
risn iN no1 v\1r× 15
Tropical estuaries are zones of very high biodiversity. Fishes form part of
this ecosystem although many may only spend part of their lives in the
estuary. Conservation and protection of tropical estuarine areas is imperative,
not only from the philosophical and aesthetic point of view, but also in order
to maintain their viability for fisheries production. The maintenance of the
ecological functioning of the estuarine ecosystem is fundamental to the
economic well being of fishers (Boisneau, 1998). They may be adversely
affected by any alterations in this functioning that affects fishes, whether or
not it is caused locally (e.g. dredging) or indirectly such as through changes in
flow (e.g. dams) or catchment use (e.g. pollutants or sediment loads). In most
tropical regions people are an integral part of the estuarine environment.
Hence any conservation planning must take their presence and activities into
account. Local communities are increasingly involved in planning and man-
agement of natural resources in tropical countries. For this reason, the value
of, for example, aquaculture developments v. retention of mangroves, is now
often questioned. There are numerous examples of local villages, particularly
in India, instigating and carrying out mangrove restoration and replanting
(Kathiresan et al., 1996). Other reasons for conserving tropical estuaries and
their fishes are international treaty obligations for maintenance of biodiversity
and perhaps more powerful, the increasing economic importance of eco-
tourism. Finally, any conservation measures should also be considered in
relation to large-scale phenomena that usually take place over relatively long
periods of time, such as climate change and sea level rise. Whether or not
these are being exacerbated by human activities may be irrelevant because they
are largely beyond control.
The many conflicts between users of estuaries and the need to maintain
their ecological viability provide a challenge to planners in many developing
as well as developed countries. Integrated planning is essential (Haq, 1997),
but in developing countries, political instability, together with lack of infra-
structure, often makes rational planning difficult and its implementation
impossible. Nevertheless, despite all these problems there is an increasing
realization that unless there is an integrated approach, involving all levels
of government, industry and a participatory approach with broad
popular support, then the future for estuaries and their fishes is bleak. In
relation to small-scale fisheries, community based management may be the
best option. In a recent review of community based management options
and systems in South East Asia, Pomeroy (1995) recognized that the under-
lying causes of over-exploitation are often social, economic or political, and
that the main focus of management should be people not fish per se.
Fisheries management in many tropical countries has followed the temperate
model of working out maximum sustainable yields and having centralized
administration (Pomeroy, 1995). The problems with this approach for the
tropics are the multispecies nature of their fisheries and the lack of consulta-
tion with the fishers who may have much of the knowledge necessary for
managing the resource. The growing realization of the need for increased
participation by resource users in fisheries management can be seen in a wide
range of policies and programmes now emerging in the South East Asian
region.
16 s. ¡. x. ni\nr×
I am most grateful to the FSBI for providing the support and opportunity to present
their paper, and to Dr Mike Elliott and colleagues for all their hospitality in Hull.
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2

. . . 

paradigms such as those related to estuarine dependence or the importance of mangroves to fisheries production have to be dealt with. In addition, no discussion of tropical estuarine fishes can ignore the human dimension, both in terms of fishing and the range of other activities that may impact on the fishes. THE ESTUARIES Many of the world’s great estuaries are in the tropics (the Amazon, Orinoco, Congo, Zambezi, Niger, Ganges and Mekong) and are all very large and receive drainage from enormous catchments. Tropical estuarine environments, however, also include tiny seasonally flowing systems of 1 to 2 km. They also encompass extensive coastal lakes and the reduced salinity estuarine waters extending along the coast in parts of South East Asia, South America and Africa. There is thus a lack of uniformity among tropical estuaries in terms of size, depth, habitats and physical regimes. These factors and many others, such as the nature of the adjacent marine and freshwater habitats, can greatly influence the fish faunas and fisheries of tropical estuaries. The world-wide extent of tropical and subtropical estuaries approximately follows the distribution of mangroves, which are the dominant intertidal vegetation in subtropical and tropical estuaries (Chapman, 1976; Duke, 1992). Mangroves generally match the 20 C isotherms in both hemispheres, suggesting that water temperature is the most significant influence. Hutchings & Saenger (1987) concluded that the presence of mangroves correlates with areas where the water temperature of the warmest month is >24 C; also that their northern and southern limits correlate reasonably well with the 16 C isotherm for the air temperature of the coldest month. Latitudinal ranges are greater on eastern continental margins than on western sides due to the presence of warm or cold currents. Tropical estuaries grade into subtropical systems beyond the tropics of Cancer and Capricorn where a winter water temperature low of c. 12 C marks their southern and northern limits (Whitfield, 1994; Ayvazian & Hyndes, 1995). Although many estuaries of the south and south east U.S.A. have been described as ‘ tropical ’ or ‘ subtropical ’ (McPherson & Miller, 1987; Powell et al., 1991), this is not really the case in a world context, with the possible exception of south Florida mangrove-lined systems. Winter water temperatures in many U.S. Gulf of Mexico estuaries fall as low as 5 C (Deegan & Thompson, 1985). To facilitate the description of those factors most relevant to their fish and fisheries, tropical estuaries can be divided into four broad categories: open estuary, estuarine coastal waters, coastal lake and blind estuary (the latter are sometimes referred to as lagoons, but this term is best avoided as it is also used in a number of other contexts, e.g. in relation to coral reefs). It must however be emphasized that any divisions are for convenience and a continuum exists.
OPEN ESTUARIES

This category includes all the larger well-known estuaries of most of the medium and larger rivers of the tropics and comes closest to the classical European and North American concept of an estuary (Elliott & McLusky, in press). These estuaries are never isolated from the sea and are subject to tidal influence with all the physical consequences of regular salinity, turbidity and

tidal prisms. the Orinoco in South America and the Mekong in Asia. mainly from trawlers.. haloclines and intertidal habitats. but estuarine conditions (defined by salinity) seldom extend beyond depths of c. which is a result of local geological history and processes. often related to the thermocline depth. They are sometimes referred to as coastal lagoons (Barnes. in the South China sea 20–50 km from the Borneo coast and in the Bay of Bengal >100 km from the mouth of the Ganges. Their faunas are mainly marine or estuarine. For example.    3 current flow changes. Important chains of coastal lakes occur along the south-east and west African coasts. and the largest of them all. This increases the pressure on fish resources and the effects of fishing may lead to changes in the fish fauna and fish community structure. The shallow nature of such tropical coastal waters and their lowered salinities and high turbidities make them at least partly estuarine in character. 1987). through an artisanal phase to fully-fledged commercial operations. often developing from a subsistence base. The cut-off point between these waters and true marine conditions may best be defined by the fauna (Pauly. to supply the demand from burgeoning cities. in the Gulf of Mexico and in northern South America. such as those of the Zambezi and Niger in Africa. What makes these systems unique is their relatively large surface area. BLIND ESTUARIES The most useful definition of blind or temporarily closed estuaries is that of Day (1981) who described them in the following terms: ‘ These are estuaries that . 1985) rather than by physical factors. The range of habitats is also determined by this connection as well as by the bathymetry. including ports and harbours with their associated industrial and city developments and consequent effluent problems. Fishery activities have increased in many of these estuaries. 1980). The form and constancy of this connection largely determines the prevailing salinities and the nature of the fish fauna. Extensive deltas occur at the mouths of many of the larger estuaries. particularly as regards their fish faunas. and most coastal lakes have some form of connection to the sea. COASTAL LAKES Many lacustrine bodies of water behind tropical shorelines on most continents are estuarine in character. ESTUARINE COASTAL WATERS Nowhere is the problem of definitions and boundaries of estuaries and estuarine fish faunas better illustrated than in tropical coastal waters. in India. Most estuarine coastal waters are subject to heavy fishing pressure. Many have been extensively modified by man and most have important fisheries. 10 km from shore (Blaber et al. Many of the larger open estuaries grade almost imperceptibly into the adjacent sea. The size and depth of this sort of estuary has made them a focus of human activity. the Ganges (Meghna) in India and Bangladesh. 1990). The effects of the discharge from the Amazon are felt up to 400 km from the mouth and those of the Orinoco up to 50 km away from the mouth. but the term lake is used to avoid confusion with small lagoons found at the mouths of blind estuaries. off Guyana these waters extend 40 km from shore (Lowe-McConnell. recreational and conservation value. 15–20 m. in the Gulf of Carpentaria c.

Orinoco estuary. particularly the composition and extent .g. particularly salinity and turbidity. and also because many of the results may not be comparable due to differences in methods. 1989. with some reaching >200. In addition. it is apparent that even small ´ estuaries in the Indo-West Pacific are usually more species rich than Atlantic systems (Table I). blind estuaries are not often subject to commercial fishing pressure.. Hence comparisons of species richness between estuaries of the Indo-West Pacific and the East and West Tropical Atlantic should be viewed with caution. 1992).. THE FISHES The species richness of fishes in tropical estuaries is greatest in the Indo-West Pacific. 20 commonly taken species [ignoring rare and adventitious species (Potter et al. ’ Estuaries in this category are usually relatively small. 5 and 15 g m 2. 1961. but are usually intensively exploited by subsistence fishers. 2002)] and warm temperate ones c. but depends upon the presence of specific habitats such as seagrass beds.  are temporarily closed by a sand bar across the sea mouth. 1986. The number of fish species in subtropical and tropical estuaries is much greater than in temperate regions. In addition. the salinity will vary. Elliott & Taylor. The composition of the fish faunas results from the interplay of a whole range of factors among which the most important are: (a) Estuary size.. Lenanton & Hodgkin. 1990). deep open water channels in the larger systems favour more of the larger species. however. 1985). Because of their relatively small size. as well as the types of habitats.. 1991. the number of species in adjacent estuarine coastal areas often exceeds 300.4 . particularly carangids and sharks. 50 (Darnell. Where individual temperate estuaries may have c. Within the tropics. Comparisons of overall fish biomasses in different tropical estuaries are difficult because there are few published figures. According to the ratio between evaporation and seepage through the bar on the one hand. equatorial examples occur in South East Asia. . it is therefore a combination of estuary size and diversity of habitats that are the most important factors in relation to species diversity. Although best developed along the warm temperate and Mediterranean climate coasts of southern Australia and South Africa (Potter et al. Elliott & Hemingway. Additionally. Freshwater enters from the river and the circulation is dependent on the residual river current and the stress of the wind on the water surface. At such times there is no tidal range and thus no tidal currents. Taking into account the large size of some Atlantic systems (e. they do extend into the sub-tropics and tropics of both Africa and India. The scarce data (Table II) suggest. These values are an order of magnitude higher than those of tropical non-estuarine seagrass areas in shallow waters (Blaber et al. both in length and catchment and are characteristic of areas of low or highly seasonal rainfall. depth and physical regimes. . but larger systems usually have more species than smaller ones. most medium to large subtropical and tropical estuarine areas have at least 100. however that fish biomass is not usually >30 g m 2 and is most commonly between c. larger systems usually have a greater diversity of habitats. and freshwater inflow plus precipitation on the other. Terminos and Lagos Lagoons). in addition to a higher number of occasional visitors. Pomfret et al.

coastal lake) in the four major zoogeographic regions (modified from Blaber. blind. (b) the nature and depth of adjacent marine waters and to a much lesser extent. both spatial and temporal. It is also important to emphasize that the number of species recorded in any system is a result of the extent of sampling. both in terms of latitude and in relation to marine features such as ocean currents. Malaysia E. fresh waters. Numbers of species recorded from different types of estuaries (O. . CL. 2000) System Indo-west Pacific Alligator Creek Trinity Embley Vellar Coleroon Chilka Hooghly Chuwei Ponggol Matang Kretam Kechil Purari Pagbilao Solomon Islands Morrumbene Tudor Creek Gazi Bay Kosi St Lucia Mhlanga East Atlantic Lagos Fatala ´ Ebrie Niger West Atlantic Guaratuba Itamaraca Orinoco Terminos Cienaga Grande Tortuguero Sinnamary East Pacific Rio Claro Guerrero Lakes Country Type and size Number of species Australia Australia Australia India India India Taiwan Singapore W. B. of mangroves.    5 T I. **Sum of species for 10 small coastal lakes. canyons and reefs. open. Malaysia New Guinea Philippines Solomon Islands* Mocambique Kenya Kenya South Africa South Africa South Africa Nigeria Guinea Ivory Coast Nigeria Brazil Brazil Venezuela Mexico Colombia Costa Rica French Guiana Costa Rica Mexico** O small O medium O large O large CL large O large O small O medium O medium O small O large O medium O small O medium O small CL small CL large CL large B Small CL large O medium CL large O large CL medium O large O large CL large CL large O small O medium O small CL small 150 91 197 195 152 172 30 78 117 44 140 128 136* 113 83 128 163 110 47 79 102 153 52 61 81 87 122 114 70 83 22 105 *Incorporates 13 small estuaries (no one system more than 50 species). (c) the geographical location of the estuary.

Florida Mexico Mexico Tropical West Atlantic Tropical East Pacific Matang Alligator Creek Embley Albatross Bay (inshore) Albatross Bay (offshore) Small estuaries Mangrove estuaries Terminos Lagoon ´ Mangrove-lined canal 0·25–3·1 2·5–29·0 5·0–16·0 5·0 12·0–30·0 11·6 15·0 0·4–3·4 7·9–12·5 Sasekumar et al. (1994) Robertson & Duke (1990) Blaber et al. (1990) Blaber & Milton (1990) Thayer et al. (1985) ˜ Warburton (1978) . (1987) Yanez-Arancibia et al. Biomasses of fishes from various subtropical and tropical estuaries (modified from Blaber.S.T II.A. 2000) Estuary or habitat Fish biomass (g m 2) Reference Country Faunal region Malaysia Australia Indo-West Pacific Solomon Islands U.. (1995) Blaber et al. (1989) Blaber et al.

g. that the division of this category from that of (a) is often difficult in the tropics. 1998) with taxonomic. 1967. McHugh.g. very euryhaline category. because many of the species that fall into the marine migrant category in the sub-tropics. Gilchristella aestuaria (Gilchrist) in Africa). both diurnally and seasonally. 1997. as well as most Leiognathidae and Haemulidae]. physiological and ecological groupings based on attributes such as salinity tolerance. Gunter. Perkins. several species of Coilia in South East Asia). engraulids (e.    7 The classification of estuarine fishes has been almost as numerous as the classification of estuaries (Day. ambassids. Sciaenidae. some species of Ariidae). Salinity tolerance may play a role in the distribution of fishes in tropical estuaries.. particularly in West Africa. The following classification scheme is similar. 1998). based mainly on how they utilize estuaries and where they spawn. Carangidae and Polynemidae) or only as adults (e. Gunter (1967) considered the lower salinity of estuarine waters to be the outstanding difference between these waters and sea water. Whitfield. For this reason. form part of the ‘ estuarine ’ category in the tropics. While this phenomenon has been amply demonstrated for warm temperate and temperate areas it is not at all clear in the subtropics and tropics. Salinity is only one of an array of factors that determine which species are found in any one subtropical or tropical estuary. a grouping of tropical estuarine fishes based mainly on their salinity tolerances is not very useful or practical. usually characterized as marine spawners. Polynemidae and Haemulidae that spawn in estuaries. This is a large group in subtropical and tropical estuaries where they may occur both as juveniles and adults [e.g. and include many species of Clupeidae. (b) Marine migrants from the sea. Blaber. . however. South East Asia and tropical South America. especially as there has been little experimental work conducted on the salinity tolerances of most species.g. or only as juveniles (e. some species of Mugilidae. Whitfield. feeding and migratory habits. a considerable degree of euryhalinity is a pre-condition and prerequisite for their inhabitants. 1974. gobies (hundreds of species). but gives increased importance to the ‘ estuarine ’ category in the tropics due to recent advances in knowledge of spawning behaviour (Blaber. the circum-tropical Mugil cephalus L. It is important to note. 1967. breeding. Based on their occurrence in low salinity waters almost all fit into a similar. Lates calcarifer (Bloch) of South East Asia and Australia. Sub-tropics: they represent a small but significant part of the estuarine fish fauna and include a number of clupeids (e. Mugilidae. 2000): (a) Estuarine species that can complete their entire life cycle in the estuary. Most research on estuarine fishes has been in temperate and warm temperate regions of Europe and North America where salinity has long been regarded as a key factor regulating the composition of estuarine fish faunas. 1981. 1951. Tropics: they represent a large part of the estuarine fish fauna. as well as for many species in marine areas that suffer seasonal reductions or fluctuations in salinity. Ariidae. 1968. atherinids and syngnathids. Green.g. but as these systems often undergo very great fluctuations in salinity (often from almost 0 to 35). This has led to a number of classification systems for subtropical and tropical estuarine fishes (Day.

A search of the Aquatic Science and Fisheries Abstracts . catadromous species are rare in the tropics (d) Freshwater migrants that move varying distances down estuaries. The dominant taxa in each region are broadly similar. The fish communities of subtropical and tropical estuaries of all four major tropical zoogeographic regions (Indo-West Pacific. this is not surprising and perhaps the term ‘ marine ’ for many of these species is a misnomer. This group is important in temperate estuaries. and secondly under pressure. but there are some interesting contrasts. In the Indo-West Pacific. but much less so elsewhere. Both these problems require detailed knowledge of the ecology of tropical estuarine fishes. Sciaenidae are one of the dominant families. West Atlantic and East Pacific) have many common characteristics. poeciliids and characids penetrate estuarine waters. eleotrids). nearly all of which are in cold or temperate regions. THE CHALLENGES THE STATUS OF TROPICAL ESTUARINE FISH RESEARCH While it is still true that most estuarine research is conducted in industrialized developed countries. It is noteworthy. where declining catches. with more than half the number of species as well as the number of individuals. they are dominated by fishes of marine origin. environmental degradation and increasing human populations are jeopardizing food security. East Atlantic. The freshwater cichlid Oreochromis mossambicus Peters and the archerfishes (Toxotidae) are well known African and South East Asian examples. there are few estuarine coastal waters and sciaenids. Given the ‘ estuarization ’ of many tropical coastal areas. but includes the commercially important tropical shads Tenualosa ilisha (Hamilton) and Tenualosa toli (Valenciennes) of south Asia and Sarawak respectively. for many of them do not occur outside of estuarine coastal areas. for conservation and maintenance of biodiversity. but usually return to fresh water to breed.8 . largely from the developed world. Except for the sciaenids.  (c) Anadromous species that breed in fresh water and on their way to and from their spawning grounds spend time in estuaries. . and although present they are not a dominant component of the fauna. In East Africa for example. In almost all cases. but is a rare category in subtropical and tropical estuaries. particularly their relationships with the environment and the extent to which they are dependent on estuaries or adjacent habitats for survival. In the tropics this group is best represented in South and Central American estuaries where a number of pimelodid catfish. being contributed by either fully estuarine species or marine migrants. however. Similarly. sciaenids are important in the equatorial regions of South East Asia. This is being driven by two imperatives: firstly that of the plight of inshore fisheries in developing countries. .g. that within the Indo-West Pacific. there has been an upsurge in tropical estuarine fish research in the last 5 years. In all regions except the Indo-West Pacific. the other dominant families are broadly comparable across all regions. sometimes to spawn (e. clupeids and engraulids are much more diverse and numerous in the estuarine coastal waters of equatorial South East Asia than in other areas. This pattern may be connected with the amount of rainfall and the degree of estuarization of coastal waters.

such as the Norman in northern . and consequently the very real decline in numbers of fish taxonomists.. BIOLOGICAL Recent advances in the knowledge of tropical estuarine fish ecology cover most aspects of the science. 1987). 1999) and Colombia (Rueda & Santos-Martinez. 1800 publications dealing with fishes in temperate estuaries. in press). A more worrying aspect is the overall decline in funding throughout the world for taxonomic research. throughout South East Asia. This is particularly the case in South and West Africa. Many are correlated with one another. Current flow in tropical estuaries. where increasing affluence and levels of education have added to many both national and international research efforts. Turbidity has been shown to be an important determinant of species composition in estuaries (Blaber & Blaber. Cyrus & Blaber. 1999). Although a number of very useful new field guides are now available (Allen. the large number of species and lack of adequate field keys frequently hamper research. salinity. Unfortunately. In terms of hydrology. has been debated for a long time. 1999). 1997). the decline in university teaching of taxonomy. one of the major obstacles to thorough ecological research still remains the difficulties associated with the correct taxonomic identification of tropical marine and estuarine fishes. much of them very relevant to understanding ecological relationships in relation to various forms of exploitation and environmental changes. South America. but the effects of current speed are not well understood. 1999) and ways of quantifying the relationship between mangroves and fisheries production are being seriously examined and studied (Baran. where syntheses of much of the previous research are being produced (Whitfield. PHYSICAL One of the most difficult challenges is to separate and evaluate the influences of each a suite of physical parameters influencing estuarine fishes. Other important practical biological problems. The role of the mangrove environment. where the results of recent research in Brazil (Chaves & Bouchereau. all are important. For example. Detailed data on the feeding and reproduction of many species are becoming available. and to a lesser extent. Following some major research efforts. particularly in developing countries. primarily as a nursery for juvenile fishes and prawns. Albaret. whereas there were only 600 for tropical estuarine fishes. 1999) are appearing. the discovery that several of the tropical clupeid shads (genus Tenualosa) are protandrous hermaphrodites has profound implications for both their fisheries management and conservation (Blaber et al. 1980. A great deal more basic data on species composition and diversity of systems are now available. accurate sampling and statistical analysis. a considerable increase in productivity from the tropics.    9 database from 1978 to the end of 1995 revealed c. current speed and turbidity. and new FAO guides are appearing. For the period from 1995 until the present the figures are 630 for temperate estuarine fishes and 338 for tropical estuarine fishes. include appropriate sampling design. 1998. particularly in developing countries where scientists seldom have access to good library facilities. the complex functional relationships are now becoming evident (Chong & Choo.

all may form part of the overall tropical estuarine community. reports to aid or funding organizations. populations. Both are also correlated with highly variable salinities and high turbidities. As stated by Blaber (2000). Here. usually muddy and hot environments with many biting insects. . ‘ hippos ’ and of course fishes. Much of the research on fishes in tropical estuaries is done in developing countries. turbidity and salinity. has more species (197) than either Groote Eylandt (179) or the relatively homogeneous Norman Estuary (100) which has a narrow band of fringing mangroves. annelid worms. comparing sites of similar size. prawns. communities or certain life history phases of fishes that occur in estuaries are dependent on the estuarine . 1987). has meant that biologists have tended to favour projects in more appealing areas. temperature and salinity have no effect on fish community structure. usually for important fisheries purposes. lunar phase and time of day or night. often with functional ecological links. Most of the variation in species composition is correlated with tidal range. particularly in mangroves (Thayer et al. Further evidence of the relative importance and interrelationships of physical factors comes from work in the Sarodrano mangroves. but the variable combinations produce communities characteristic of particular conditions. this is reflected by the greater number of species found in the complex forests of the Embley than in the more depauperate Norman Estuary. tidal range. such as coral reefs. including a wide variety of mangroves. 1994). results largely from tidal range and freshwater inflow. birds. The relative importance of each factor varies from estuary to estuary. and not necessarily for fundamental scientific reasons.10 . and the turbidity gradient is constant throughout the year and does not cause any seasonal variations in the distribution of juveniles. habitat diversity and structure are important in relation to species composition. often linked to poverty alleviation.  Australia (Blaber et al. Many of the results are only to be found in the ‘ grey literature ’ of annual reports. POPULAR PARADIGMS—ESTUARINE DEPENDENCE Whether or not particular species. unpublished conference papers and articles in newsletters. LOGISTIC Tropical estuaries contain a wide range of taxa that are very closely associated.. It is thus probable that the fish communities of estuarine and inshore waters of the tropical Indo-West Pacific are at least partly determined by the interrelationships between current speed.. In addition. the difficulties of working in these often inhospitable. whether or not they are important. the Embley Estuary.. crocodiles. as well as by the structure of the habitat. within the Gulf of Carpentaria in northern Australia. Resources and facilities in such areas are usually limited or sometimes non-existent and publication of results. Unfortunately. 1997). may not occur. with a broad range of habitats. The influence of the structural heterogeneity of each habitat is important. in a semi-arid area of Madagascar (Laroche et al. such data are often important and may be the only information on a particular area or species and it is vital to draw fully on this literature to support and extend results from the primary literature. .

for example. Briggs (1974) proposed that the tropical Indo-Malayan region was the centre of speciation among Indo-West Pacific shallow water fishes. It is important to note. 5%). Unfortunately there is little agreement in general terms about the validity of ‘ estuarine dependence ’. species utilizing the estuary as a nursery make up 35 to 72% of all species. In these non-tropical areas many marine species have been shown to be dependent on the estuarine environment during their juvenile phase (Day et al. 1967. 28%). Thryssa and Elops are common in estuarine coastal waters throughout South and South East Asia. 1982). Blaber (1981) and Blaber & Blaber (1980) postulated that most of the estuarine fish fauna of the sub-tropics and tropics is not ‘ estuarine ’ per se. and Australia. often with variable salinity.S. such as Gerres. For example in the estuaries of the Louisiana deltaic plain. 1985). Deegan & Thompson. turbid areas. form what Blaber (1997) termed the ‘ core area ’ for the majority of Indo-West Pacific estuarine species. however. from coral reefs to seagrasses to mangroves and estuaries. marine species (occur in estuaries in small numbers as occasional visitors. where vast areas of estuarine coastal waters in South and South East Asia. McHugh. 3%). 1981.S. Many of the taxa considered largely estuarine dependent in.. euryhaline freshwater species (may breed in estuaries. but are confined mainly to estuaries in southern Africa.A. West Africa and northern South America. and as discussed earlier.S. are not strictly estuarine dependent in the tropics. and in southern Africa. refugia or spawning. due mainly to parochialism. Whether the greater number of species in this region is due to enhanced . 1985). with its unparalleled variety of habitats. and the all too ready acceptance by workers everywhere of the paradigm developed for fishes of the estuaries of the south eastern U. The concept of an estuarine dependent phase in the life ´˜ cycle of coastal fishes arose primarily from studies of temperate and warm temperate estuaries in the U.A. Whitfield (1994) divided the 142 estuarine fishes into five categories in relation to estuarine dependence: estuarine species (species that breed in estuaries. Day & Yanez-Arancibia. They concluded that only in Brazil and South Africa. The region with the greatest diversity of coastal fish species is the tropical Indo-West Pacific. South Africa. and parts of the U. that this classification includes a mixture of temperate and warm temperate species as well as those of subtropical South East Africa. Areas with such characteristics occur over very large areas of the sea in South and South East Asia. in systems that are marginally subtropical. could estuarine dependence really be demonstrated. south to parts of northern Australia. and usually over 90% of individuals (Deegan & Thompson. The degree of estuarine dependence among fishes of the sub-tropics and tropics was questioned by Longhurst & Pauly (1987) who analysed studies from 20 subtropical and tropical areas.A. but is a fauna characteristic of shallow. 43%). (Gunter. 21%). euryhaline marine species (species that breed in the sea and juveniles occur in estuaries.    11 environment for feeding. Therefore according to this classification 71% of the species are completely or partially dependent on estuaries. as opposed to the sea or even fresh water. 1985). 1967. and obligate catadromous species (species that use estuaries in transit between the sea and fresh water. In southern Africa. has given rise to much discussion and speculation (Hedgpeth. many of the subtropical species included in this classification are at the southern limits of their distribution.

For example. a causal link has not been established experimentally. Certainly. east Australian and south Pacific shores where shallow.12 . Nevertheless. Almost all species in the estuaries of northern Australia are also common throughout South and South East Asia. Many more species spawn in tropical estuarine areas. habitats. two thirds of the species in the St Lucia coastal lake system of KwaZulu-Natal. Such marginal areas are restricted to the relatively small areas of estuaries along south-east African. including particularly the Sciaenidae. 1974). The situation with regard to fishes is less clear. (1985) showed a positive correlation between commercial fish catches and mangrove area in the Gulf of Mexico. and fish community structure have not been worked out fully. 1977). 1998). Interesting comparative information is available for some species of penaeid prawns that like many fish species. turbid and variable salinity conditions give way to coral reefs and clearer waters. De Graaf & Xuan (1998) demonstrated a similar relationship. and Barbier & Strand (1998) determined the effects of changes in mangrove area in the Laguna de Terminos (Gulf of Mexico) on the production and value of prawn harvests in Campeche from 1980 to 1990. particularly where physical conditions change and mangrove habitats are not dominant (Blaber. where most species remain in similar (large and widespread) habitats for their whole life. Resolving the question of the importance of ‘ estuarine dependence ’ for tropical fishes is hampered because the interrelationships between environmental factors. but the question of total estuarine dependence may only be relevant in estuaries outside the ‘ core areas ’. Engraulididae and Clupeidae. 2000). provides good evidence that there is a correlation between commercial offshore prawn catches and the total area of adjacent mangroves. 1985) and the Philippines (Primavera. In southern Vietnam. The majority of species found in tropical estuaries are probably not estuarine dependent.  speciation rates or reduced extinction rates remains a subject of controversy (Briggs.. lies the main research challenge in relation to ‘ estuarine dependence ’ in tropical fishes. and led to Robertson & Blaber (1992) concluding. Indonesia (Martosubroto & Naamin. than they do in temperate or warm temperate estuaries. the similarity of tropical estuarine fish faunas across the great breadth of the Indo-West Pacific is striking. For most families there is also a gradual reduction in species diversity away from the ‘ core area ’. Yanez-Arancibia et al. Research in the Gulf of Mexico (Turner. Here perhaps. . Australia (Staples et al. 1977). POPULAR PARADIGMS—MANGROVES AND FISHERIES PRODUCTION The degree to which the size and productivity of coastal and offshore fisheries in the tropics is dependent on the extent of healthy nursery or feeding areas in estuaries or mangroves has given rise to much debate. that in spite of apparent evidence of a correlation between mangrove area and commercial fish catches. South Africa and 71% of species in Madagascar estuaries occur in South East Asia. It has less relevance where there is extensive estuarization (Longhurst & Pauly. 1987) of the continental shelf. In the . . have a juvenile phase that lives in estuaries and mangroves. at a species level. Pauly & Ingles (1986) concluded that most of the variance in the catches of penaeids could be explained by a combination of mangrove area and latitude.

it was calculated that one hectare of mangrove forest supports a marine catch of 450 kg year 1. and finally at meaningful economic evaluations. contributing to the relationships. Only by using accurate production figures and a suite of ecological data for each species assemblage.    13 latter study. as stated by Baran (1999) and Baran & Hambrey (1999). Tropical estuaries are among the most exploited ecosystems in the world. the nursery function of mangroves is highly variable at local. 75% of nursery function from 50% of original area (Kapetsky. where any increases in fishing effort are concentrated on juveniles this may affect adult (fishable) stocks. including scientific. sociological (including cultural) and economic. Unfortunately. Hence. physical and biological data offers the only hope for arriving at both an understanding of the fisheries and mangrove relationship. perhaps much of the functional value could be retained. These authors conclude that the integration of economic. As the focal . It is possible that the mangrove forests fringing deeper water contain much of the functionality compared with the more inland shallower. area of shallow coastal water. Unfortunately. Baran & Hambrey (1999) discussed the economic value of the fisheries function of mangroves and indicate that most published monetary values are misleading because they use aggregated fisheries production and uniformity of mangrove nursery function. all the studies to date suffer from problems of auto-correlation. It requires the integration of a number of disciplines and inputs. intertidal areas (Vance et al. but it must again be reiterated that these are correlations and that causal links have not been established experimentally. taking account of the activity of people in estuaries is perhaps the hardest challenge of all. and deeper fringing areas left intact. The few studies that have quantified relationships between mangroves and coastal resources (mainly penaeid prawns) were summarized by Baran (1999) (Table III). if much of the loss of mangroves could be confined to the inland side. There is also no general agreement about whether any such relationship between mangrove area and fisheries production is linear. and improved fishing technology. not only with massive reductions of mangrove area. 1985). as stated by Robertson & Blaber (1992) and Baran & Hambrey (1999). with many factors other than just mangrove area. 1999). but also with great increases in fishing effort. In fact. It has been suggested that much of the functional value of mangroves might be retained from a smaller area of mangroves. Overall declines in fish catches in many parts of the tropics coincide. THE HUMAN DIMENSION From a fisheries research or management perspective. Also. intertidal area and food availability.. such as river discharge. Hence the ability to use any time-series data to quantify fisheries-mangrove relationships is compromised. for example. especially for biologists. The calculation of any historical relationships between fisheries production and mangrove area are complicated by changes in both fishing effort and fishing methods. 1996). would it be possible to provide meaningful economic values (Baran & Hambrey. in recent important reviews of the subject. regional and global scales. it is these deeper fringing areas of mangroves adjacent to the sea that have been most attractive and suitable for aquaculture pond development throughout the tropics.

T III.A. 1999) Formula Lny=0·496Lnx+6·07 y=1·96x y=0·1128x+5·473 4·39 Coastal marshes in km2 x function y function Fish capture (t) r2 n 0·48 10 0·92 7 0·79 — Log10 of penaeid catch (t) 0·66 17 Catch of Penaeus merguiensis (t) 0·58 6 Source and area Yanez-Arancibia et al. Martosubroto & Naamin (1977)— Indonesia Paw & Chua (1989)—South East Asia Staples et al. AM. Maximum sustainable yield of penaeids. (1985)—Australia Pauly & Ingles (1986)—tropics % of saline vegetation in a Percentage of brown shrimps hydrological unit Mangrove area ( 10 000 ha) Penaeid production ( 1000 t) y=0·8648x+0·0991 Log10 of mangrove area y=1·074x+218·3 Mangrove shoreline in km Log10MSY=0·4875log10 AM 0·0212L+2·41 MSY. (1985)— ˜ Mexico Turner (1977)—U. L. . degrees latitude. Relationships between mangroves and fisheries production that have been quantified (after Baran. area of mangroves.S.

A. Production of fish in different types of tropical estuaries (modified from Blaber. 1999. El Salvador Philippines South Africa Ivory Coast India Ghana Malagasy Colombia Mexico Venezuela Estuary Hooghly-Matlah Vellar-Coloroon Larut-Matang Texas bays Jiquilisco San Miguel Bay Kosi system Ebrie Lagoon u ´ Lake Chilka Lake Pulicat Mandapam Lagoon Sakumo Lagoon Pangalanes Lagoon Cienaga Grande Caimanero Lagoon Terminos lagoon Tamiahua Lagoon Lake Maracaibo Tacarigua Lagoon Type Open estuary system Open estuary system Open estuary system Estuarine coastal waters Estuarine coastal waters Estuarine coastal waters Coastal lakes and open estuary Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake Coastal lake 2 T km 11·4 11·1 38·64* 12·1 1·7 23·8** 1·0 16·0 3·7 2·6 5·6 15·0 3·7 12·0 34·5 20·0 4·7 1·9 11·0 2 *Includes penaeid prawn catches and not coastal waters. Allied to the fishing industry is that of aquaculture. . or removal of the habitat and the indirect effects of changing fish community structure by selectively removing some species. Nevertheless. Most large tropical or subtropical estuaries have port cities associated with them that usually have major industries that pour their effluents into the estuary. Industries bring population increases and growth of cities. The development of large-scale aquaculture ventures in tropical estuaries has usually been at the expense of mangroves and ultimately wild fisheries production. they have provided quiet and sheltered waters for harbours. 2000). 2000) Country India Malaysia U. The effects of fishing itself is a subject that is receiving much attention throughout the world (Hall. Such effects in estuaries include not just overfishing. and historically gave the easiest or only access to the interior for trade or settlement. but also alterations to.    15 T IV. which discharge their sewage into the estuary.. Blaber et al. 1997). For peoples living in the interior they have acted as a route to the sea for trade. Hence the overriding need to reconcile conflicts between uses and users. most tropical estuaries are still vitally important as a source of fishes to the local people and fishing plays an important role in the economy (Table IV). fishing and migration. which together with discharges from shipping. Although they may have the highest economic value per hectare of any aquatic environment (Costanza. they also have some of the most intractable problems. have resulted in the lower reaches of many estuaries becoming badly polluted.S. point for much human activity at the meeting place of sea and river. **Probably an overestimate as trawlable biomass only 2·13 t km .

any conservation measures should also be considered in relation to large-scale phenomena that usually take place over relatively long periods of time. is now often questioned. pollutants or sediment loads). 1997). and that the main focus of management should be people not fish per se. instigating and carrying out mangrove restoration and replanting (Kathiresan et al. aquaculture developments v. Conservation and protection of tropical estuarine areas is imperative. They may be adversely affected by any alterations in this functioning that affects fishes. For this reason. retention of mangroves. In most tropical regions people are an integral part of the estuarine environment. but also in order to maintain their viability for fisheries production.16 .g.g. dredging) or indirectly such as through changes in flow (e. community based management may be the best option. often makes rational planning difficult and its implementation impossible. involving all levels of government. 1996). the increasing economic importance of ecotourism. together with lack of infrastructure. Whether or not these are being exacerbated by human activities may be irrelevant because they are largely beyond control. not only from the philosophical and aesthetic point of view. the value of. dams) or catchment use (e. The problems with this approach for the tropics are the multispecies nature of their fisheries and the lack of consultation with the fishers who may have much of the knowledge necessary for managing the resource. Finally. such as climate change and sea level rise. Nevertheless. The maintenance of the ecological functioning of the estuarine ecosystem is fundamental to the economic well being of fishers (Boisneau. The many conflicts between users of estuaries and the need to maintain their ecological viability provide a challenge to planners in many developing as well as developed countries.. economic or political. Fisheries management in many tropical countries has followed the temperate model of working out maximum sustainable yields and having centralized administration (Pomeroy. 1998). particularly in India. industry and a participatory approach with broad popular support.g. despite all these problems there is an increasing realization that unless there is an integrated approach. . In a recent review of community based management options and systems in South East Asia. then the future for estuaries and their fishes is bleak. There are numerous examples of local villages. political instability. 1995). Pomeroy (1995) recognized that the underlying causes of over-exploitation are often social. Other reasons for conserving tropical estuaries and their fishes are international treaty obligations for maintenance of biodiversity and perhaps more powerful. for example.  Tropical estuaries are zones of very high biodiversity. The growing realization of the need for increased participation by resource users in fisheries management can be seen in a wide range of policies and programmes now emerging in the South East Asian region. . Integrated planning is essential (Haq. . In relation to small-scale fisheries. Fishes form part of this ecosystem although many may only spend part of their lives in the estuary. whether or not it is caused locally (e. Local communities are increasingly involved in planning and management of natural resources in tropical countries. Hence any conservation planning must take their presence and activities into account. but in developing countries.

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