Journal of Fish Biology (2002) 61, (Supplement A), 120

doi:10.1006/jfbi.2002.2063, available online at http://www.idealibrary.com on

Fish in hot water : the challenges facing sh and sheries
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 worlds 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 shers, industrialists, shippers, farmers, conservationists, sports
enthusiasts and biologists overlap to such an extent. Quite apart from the possible eects of all
these activities, the shes 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 shes, all may form
part of the overall community, often with functional ecological links. Unfortunately, the
diculties 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 sh 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 shes,
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: sheries; tropical estuaries; research; sh; biodiversity.
INTRODUCTION
This review considers shes 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
shes! 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 briey describe the great diversity of their
shes; the similarities between the sh 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 shes, 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
00221112/02/61A001+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 sheries production have to be dealt with. In addition, no
discussion of tropical estuarine shes can ignore the human dimension, both in
terms of shing and the range of other activities that may impact on the shes.
THE ESTUARIES
Many of the worlds 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 owing 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
inuence the sh faunas and sheries 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 signicant inuence. 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 (Whiteld, 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 sh and
sheries, 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
inuence with all the physical consequences of regular salinity, turbidity and
2 s. . x. ni\nr
current ow 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 euent problems. Fishery activities have
increased in many of these estuaries, often developing from a subsistence base,
through an artisanal phase to fully-edged commercial operations, to supply the
demand from burgeoning cities. This increases the pressure on sh resources and
the eects of shing may lead to changes in the sh fauna and sh community
structure.
ESTUARINE COASTAL WATERS
Nowhere is the problem of denitions and boundaries of estuaries and
estuarine sh faunas better illustrated than in tropical coastal waters. Many of
the larger open estuaries grade almost imperceptibly into the adjacent sea. The
eects 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 sh
faunas. The cut-o point between these waters and true marine conditions
may best be dened by the fauna (Pauly, 1985) rather than by physical factors,
but estuarine conditions (dened by salinity) seldom extend beyond depths of
c. 1520 m, often related to the thermocline depth. For example, o 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
2050 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 shing
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 sh 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 modied by man and
most have important sheries, 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 denition 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 inow 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 shing pressure, but are usually intensively exploited by
subsistence shers.
THE FISHES
The species richness of shes in tropical estuaries is greatest in the Indo-West
Pacic, 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
Pacic 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 Pacic are usually more species rich than Atlantic
systems (Table I). Comparisons of overall sh biomasses in dierent tropical
estuaries are dicult because there are few published gures, and also because
many of the results may not be comparable due to dierences in methods. The
scarce data (Table II) suggest, however that sh 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 sh 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
specic habitats such as seagrass beds. The composition of the sh 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 dierent types of estuaries (O, open; B, blind;
CL, coastal lake) in the four major zoogeographic regions (modied from Blaber, 2000)
System Country Type and size
Number of
species
Indo-west Pacic
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 Pacic
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.
risn iN no1 v\1r 5
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The classication of estuarine shes has been almost as numerous as the
classication of estuaries (Day, 1951; Gunter, 1967; McHugh, 1967; Green,
1968; Perkins, 1974; Whiteld, 1998) with taxonomic, physiological and
ecological groupings based on attributes such as salinity tolerance, breeding,
feeding and migratory habits. Most research on estuarine shes 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 sh faunas. Gunter (1967) considered the lower salinity of estuarine
waters to be the outstanding dierence 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 shes in tropical estuaries, but as these systems often undergo
very great uctuations 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
suer seasonal reductions or uctuations in salinity.
For this reason, a grouping of tropical estuarine shes 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 t into a similar, very
euryhaline category. This has led to a number of classication systems for
subtropical and tropical estuarine shes (Day, 1981; Blaber, 1997; Whiteld,
1998), based mainly on how they utilize estuaries and where they spawn. The
following classication 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 signicant part of the estuarine sh
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 sh
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
dicult 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 catsh, poeciliids and
characids penetrate estuarine waters. The freshwater cichlid Oreochromis
mossambicus Peters and the archershes (Toxotidae) are well known
African and South East Asian examples.
The sh communities of subtropical and tropical estuaries of all four major
tropical zoogeographic regions (Indo-West Pacic, East Atlantic, West Atlantic
and East Pacic) have many common characteristics. In almost all cases, they
are dominated by shes 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 Pacic,
Sciaenidae are one of the dominant families. In the Indo-West Pacic, 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 Pacic, 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 sh research in the last 5 years. This is
being driven by two imperatives: rstly that of the plight of inshore sheries 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 shes, 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
shes in temperate estuaries, whereas there were only 600 for tropical estuarine
shes. For the period from 1995 until the present the gures are 630 for
temperate estuarine shes and 338 for tropical estuarine shes, a considerable
increase in productivity from the tropics.
BIOLOGICAL
Recent advances in the knowledge of tropical estuarine sh 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 (Whiteld, 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 auence and levels of education have added to many both national
and international research eorts. 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 sheries management and conservation
(Blaber et al., in press). The role of the mangrove environment, primarily as a
nursery for juvenile shes and prawns, has been debated for a long time.
Following some major research eorts, the complex functional relationships are
now becoming evident (Chong & Choo, 1999) and ways of quantifying the
relationship between mangroves and sheries production are being seriously
examined and studied (Baran, 1999).
Unfortunately, one of the major obstacles to thorough ecological research still
remains the diculties associated with the correct taxonomic identication of
tropical marine and estuarine shes. Although a number of very useful new eld
guides are now available (Allen, 1997), and new FAO guides are appearing, the
large number of species and lack of adequate eld 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 sh
taxonomists. Other important practical biological problems, particularly in
developing countries, include appropriate sampling design, accurate sampling
and statistical analysis.
PHYSICAL
One of the most dicult challenges is to separate and evaluate the inuences
of each a suite of physical parameters inuencing estuarine shes. 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 eects of current speed are not well
understood. Current ow 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
inow. 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 eect on sh 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 inuence of the structural heterogen-
eity of each habitat is important, particularly in mangroves (Thayer et al., 1987);
this is reected 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 sh communities of estuarine and inshore waters of the tropical
Indo-West Pacic 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 shes, all may
form part of the overall tropical estuarine community, often with functional
ecological links. Unfortunately, the diculties 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 shes in tropical estuaries is done in developing
countries, usually for important sheries purposes, often linked to poverty
alleviation, and not necessarily for fundamental scientic 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 PARADIGMSESTUARINE DEPENDENCE
Whether or not particular species, populations, communities or certain life
history phases of shes 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 shes of the
estuaries of the south eastern U.S.A. (Gunter, 1967; McHugh, 1967; Day &
Yan ez-Arancibia, 1985). The concept of an estuarine dependent phase in the life
cycle of coastal shes 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 shes 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 sh 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 conned 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, Whiteld (1994) divided the 142 estuarine shes into ve
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
classication 71% of the species are completely or partially dependent on
estuaries. It is important to note, however, that this classication 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 classication
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 sh species is the tropical
Indo-West Pacic, 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 Pacic estuarine species.
Briggs (1974) proposed that the tropical Indo-Malayan region was the centre of
speciation among Indo-West Pacic shallow water shes, 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 sh faunas
across the great breadth of the Indo-West Pacic 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
Pacic 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 shes is hampered because the interrelationships between environmental
factors, habitats, and sh community structure have not been worked out
fully. Here perhaps, lies the main research challenge in relation to estuarine
dependence in tropical shes.
POPULAR PARADIGMSMANGROVES AND FISHERIES PRODUCTION
The degree to which the size and productivity of coastal and oshore sheries
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 sh
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 oshore 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 shes is less clear, and led to Robertson & Blaber
(1992) concluding, that in spite of apparent evidence of a correlation between
mangrove area and commercial sh catches, a causal link has not been
established experimentally. Certainly, Yanez-Arancibia et al. (1985) showed a
positive correlation between commercial sh catches and mangrove area in the
Gulf of Mexico, and Barbier & Strand (1998) determined the eects 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 quantied 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 suer 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 sheries 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 conned 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 sheries production
and mangrove area are complicated by changes in both shing eort and shing
methods. Overall declines in sh catches in many parts of the tropics coincide,
not only with massive reductions of mangrove area, but also with great increases
in shing eort, and improved shing technology. Also, where any increases in
shing eort are concentrated on juveniles this may aect adult (shable) stocks.
Hence the ability to use any time-series data to quantify sheries-mangrove
relationships is compromised.
Baran & Hambrey (1999) discussed the economic value of the sheries
function of mangroves and indicate that most published monetary values are
misleading because they use aggregated sheries 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 gures
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 oers the
only hope for arriving at both an understanding of the sheries and mangrove
relationship, and nally at meaningful economic evaluations.
THE HUMAN DIMENSION
From a sheries 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 scientic, 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, shing 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 euents 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
shes to the local people and shing plays an important role in the economy
(Table IV). Hence the overriding need to reconcile conicts between uses and
users. The eects of shing itself is a subject that is receiving much attention
throughout the world (Hall, 1999; Blaber et al., 2000). Such eects in estuaries
include not just overshing, but also alterations to, or removal of the habitat and
the indirect eects of changing sh community structure by selectively removing
some species. Allied to the shing 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 sheries production.
T\nir IV. Production of sh in dierent types of tropical estuaries (modied from
Blaber, 2000)
Country Estuary Type T km
2
India Hooghly-Matlah Open estuary system 114
Vellar-Coloroon Open estuary system 111
Malaysia Larut-Matang Open estuary system 3864*
U.S.A. Texas bays Estuarine coastal waters 121
El Salvador Jiquilisco Estuarine coastal waters 17
Philippines San Miguel Bay Estuarine coastal waters 238**
South Africa Kosi system Coastal lakes and open estuary 10
Ivory Coast Eubrie Lagoon Coastal lake 160
India Lake Chilka Coastal lake 37
Lake Pulicat Coastal lake 26
Mandapam Lagoon Coastal lake 56
Ghana Sakumo Lagoon Coastal lake 150
Malagasy Pangalanes Lagoon Coastal lake 37
Colombia Cienaga Grande Coastal lake 120
Mexico Caimanero Lagoon Coastal lake 345
Terminos lagoon Coastal lake 200
Tamiahua Lagoon Coastal lake 47
Venezuela Lake Maracaibo Coastal lake 19
Tacarigua Lagoon Coastal lake 110
*Includes penaeid prawn catches and not coastal waters.
**Probably an overestimate as trawlable biomass only 213 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 sheries production. The maintenance of the
ecological functioning of the estuarine ecosystem is fundamental to the
economic well being of shers (Boisneau, 1998). They may be adversely
aected by any alterations in this functioning that aects shes, whether or
not it is caused locally (e.g. dredging) or indirectly such as through changes in
ow (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 shes 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 conicts 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 dicult 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 shes is bleak. In
relation to small-scale sheries, 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 sh 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 sheries and the lack of consulta-
tion with the shers 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 sheries 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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