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BULLETIN OF MARINE SCIENCE, 80(3): 457–472, 2007

457Bulletin of Marine Science© 2007 Rosenstiel School of Marine and Atmospheric Science of the University of Miami

FIRST INTERNATIONAL SYMPOSIUM ON MANGROVES AS FISH HABITAT KEYNOTE ADDRESS

MANGROVES AND FISHES: ISSUES OF DIVERSITY, DEPENDENCE, AND DOGMA

Stephen J. M. Blaber

ABSTRACTTropical estuarine fishes are inextricably linked with mangroves, which are the

dominant vegetation of tropical and subtropical estuaries. Among the most produc-tive of aquatic areas and heavily exploited, their future may depend upon ecosystem understanding. This paper reviews diversity, dependence, and connectivity between mangroves and fisheries in the light of data from previously unstudied systems in developing countries and new approaches in developed countries. Fish diversity in mangroves varies at global, latitudinal, regional, local and habitat scales, and spe-cies composition in any one system represents the combined influences of factors operating at each of these scales. Mangrove dependence paradigms require criticalMangrove dependence paradigms require critical evaluation as new data become available and as catches and mangrove areas decline. Although it is a widely held dogma that mangroves are essential for fish populations, most evidence is circumstantial. Therefore experimental and quantitative studies are needed to support arguments that the value of retaining mangroves exceeds that of their destruction.

Mangroves are the dominant vegetation and one of the most characteristic habi-tats of tropical and subtropical estuaries. Their approximate distribution is bounded north and south of the equator by the 20 °C isotherm. The more than 80 species of mangroves form forests and stands varying in complexity from single species fringes in most subtropical estuaries to multi-species forests covering many square kilome-ters in tropical systems. Some mangrove forests in the tropics have a complex zona-tion, may contain upwards of 17 species of trees (Hutchings and Saenger, 1987), and are very dense with many prop roots. The largest mangrove forests with greatest di-versity occur in southeast Asia, northern Australia, and northeast South America.

Any discussion of fishes in tropical inshore and estuarine areas is inextricably linked with mangroves, areas of high fish species diversity and complex interrela-tionships. More than 200 species of fish can occur in any one mangrove estuary of the tropical Indo-West Pacific, with somewhat fewer, although still more than 100, in tropical East Atlantic and neotropical estuaries (Blaber, 2000). These systems are among the most productive on the planet (Costanza et al., 1997) and hence are heavily exploited by man. The present total worldwide area of mangroves has been estimated at < 150,000 km2 (FAO, 2003), but this represents only about 40% of the original cover. Many countries, such as Thailand, Philippines, and Ecuador, have lost far more than 50% of their mangroves; Singapore only has 0.5% of their mangroves remaining (Sasekumar and Wilkinson, 1994). One of the major factors in the loss of estuarine habitat in the subtropics and tropics has been the destruction of huge areas of mangroves for aquaculture, timber, and industrial developments. Most man-grove systems in the tropics are in developing countries where fisheries are heavily exploited and issues of food security usually override management and conservation considerations. There is now the realization that for sustainable management to be successful humans must be included as part of the tropical estuarine ecosystem (Fig. 1) (Blaber, 2005).

BULLETIN OF MARINE SCIENCE, VOL. 80, NO. 3, 2007458

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BLABER: MANGROVES AND FISHES—DIVERSITY, DEPENDENCE, AND DOGMA 459

Against this background there is increasing concern about the future integrity and sustainable management of mangrove systems and their associated fishes. The need to understand the interrelationships between mangroves and fishes is now greater than ever, not only for sustainable fisheries management reasons, but for overall man-agement of the exploitation or conservation of mangrove forests. Such understand-ing involves a number of concepts, particularly those concerning (1), the varying diversity of mangrove-associated fishes and the major influences causing differences among fish faunas of mangrove systems; (2), the various paradigms about estuarine or mangrove dependence of fishes and their validity; and (3), the relationships be-tween mangroves and fisheries production. Each of these three points will be re-ex-amined in this paper in the light of new data, especially from previously unstudied systems in developing countries and new approaches in developed countries.

Relationships Between Mangroves and Fish Species Diversity.—The di-versity of fishes in mangroves varies significantly at a number of scales; global, latitu-dinal, regional, local, and among habitats within individual systems. The number of species in a system represents the combined influences of factors operating at each of these scales. It is necessary to examine the role of each of these scales because the species composition in any one place is the result of their combined influences, and results in every system being unique.

At a global scale, in terms of the world’s major zoogeographic regions, the highest diversities are found in the tropical Indo-West Pacific, a huge area stretching from the coast of East Africa to the central Pacific, where upwards of 600 species have been recorded in mangrove estuaries and many individual systems may have as many as 200 species (Table 1). The high diversity decreases latitudinally away from the equatorial “core area” (sensu Blaber, 2000) in southeast Asia, but larger sub-tropical mangrove systems still contain at least 100 species. The mangroves of the Tropical East Atlantic region along the West African coast have somewhat fewer species, but are still relatively rich, with larger estuaries such as the Senegal, having more than 130 species and smaller systems such as the Fatala in Guinea about 100 species. The Tropical West Atlantic region from the Gulf of Mexico to northern South America has similar numbers of species, with most open estuaries in the equatorial region containing at least 100 species. There is some attenuation away from the equator, although species numbers are more influenced by size and type of system rather than by latitude (Table 1).

At a regional level, species composition is influenced mainly by habitat diversity, structure, and hydrology. For example, a comparison of the large Embley and Norman estuaries, both in the Gulf of Carpentaria, Australia, reveals that the former has twice as many species. The Norman has a narrow fringe of mangroves, soft muddy substrata, a large tidal range (> 4 m), extremes of salinity, and high current speeds, whereas the Embley has a broad range of habitats, including extensive mangrove forests, a variety of substrata, and less extreme hydrological conditions (Blaber et al., 1994). The pos-sible influences on species composition of extensive mangrove forests vs smaller fring-ing mangroves is illustrated by the Matang and Merbok estuaries in West Malaysia. The former has large and diverse mangrove forests covering many square kilometers, while in the latter, large tracts of mangroves have been removed and there are now only fringing stands of mangroves. The Matang system has at least 117 species while about half this number have been recorded in the Merbok (Khoo, 1989; Chong et al., 1990; Sasekumar et al., 1994; Hayase and Fadzil, 1999). Although the numbers of species in


these two systems are different, the dominant families in both are the Ariidae and Sci-aenidiae. However, the order of abundance of species in these families is quite different in the two estuaries, especially among the Sciaenidae.

Table 1. Numbers of species recorded from different types of mangrove systems (O = open, B = blind, CL = coastal lake) in the four major zoogeographic regions (modified from Blaber, 2000).

System Country Type and size No. of spp. Source

Indo-West PacificAlligator Creek Australia O small 150 Robertson and Duke (1990)Trinity Australia O medium 91 Blaber (1980)Embley Australia O large 197 Blaber et al. (1989)Vellar Coleroon India O large 195 Krishnamurthy and

Jeyaseelam (1981)Chilka India CL large 152 Jones and Sunjansingani

(1954)Chuwei Taiwan O small 30 Lin and Shao (1999)Ponggol Singapore O medium 78 Chua (1973)Matang W. Malaysia O medium 117 Sasekumar et al. (1994)Kretam Kechil E. Malaysia O small 44 Inger (1955)Purari New Guinea O large 140 Haines (1979)Pagbilao Philippines O medium 128 Pinto (1988)Solomon Islands Solomon Islands* O small 136* Blaber and Milton (1990)Morrumbene Mozambique O medium 113 Day (1974)Tudor Creek Kenya O small 83 Little et al. (1988)Kosi South Africa CL large 163 Blaber and Cyrus (1981)Mhlanga South Africa B Small 47 Harrison and Whitfield (1995)

East AtlanticLagos Nigeria CL large 79 Fagade and Olaniyan (1972)Fatala Guinea O medium 102 Baran (1995)Ebrié Ivory Coast CL large 153 Albaret (1994)Niger Nigeria O large 52 Boeseman (1963)Gambia Gambia O large 89 Baran (2000)Sine-Saloum Senegal O large 123 Baran (2000)

West AtlanticCaeté Brazil O medium 82 Barletta et al. (2005)Guaratuba Brazil CL medium 61 Chaves and Bouchereau

(1999)Itamaraca Brazil O large 81 Paranagua and Eskinazi-Leca

(1985)Orinoco Venezuela O large 87 Cervigon (1985)Ciénaga Grande Colombia CL large 114 Leon and Racedo (1985)Tortuguero Costa Rica O small 70 Gilbert and Kelso (1971)Sinnamary French Guiana O medium 83 Boujard and Rojas-Beltran

(1988)Everglades USA O large 53 Thayer et al. (1987)

East PacificRío Claro Costa Rica O small 22 Lyons and Schneider (1990)Guerrero Lakes Mexico** CL small 105 Yáñez-Arancibia (1978)* incorporates 13 small estuaries — no one system more than 50 species.** Sum of species for 10 small coastal lakes.


At a more local level, it was shown that various mangrove habitats in Biscayne Bay Florida harbor different fish assemblages depending upon a suite of factors, the most important of which are (1) proximity to offshore reef habitats, (2) the salin-ity regime along the shoreline, and (3) the water depths within the mangrove for-est interior (Serafy et al., 2003). Fish assemblages also vary in small Solomon Island estuaries (Blaber and Milton, 1990), but here the major influences are (1) the species of mangrove (i.e., type of structure), (2) whether the channels are blocked or choked by fallen mangrove branches, and (3) the type of substratum (ranging from soft mud to hard sand and rock). In the Pagbilao mangroves in the Philippines, Rönnbäck et al. (1999) found that fish species composition and abundance were influenced by (1) the dominant mangrove species and the structural complexity of the root system, (2) proximity to open water habitat, and (3) water depth.

All of these examples serve to illustrate that the composition of mangrove fish as-semblages is determined by an interplay of factors that include structural diversity of the habitat, hydrological features of current speed, tidal range, turbidity and salinity, and the nature of adjacent waters. The fish assemblage in any one mangrove system is the result of the combined effects of all of these factors and the relative importance of each.

One factor in the interrelationship between mangroves and fishes that appears to have received little attention is the age of a mangrove. Most mangrove forests are dy-namic and undergo constant change with various successions of tree species and tree sizes that are relatively well understood in a growing mangrove forest (Hutchings and Saenger, 1987). In the Matang mangroves of Malaysia, density and biomass of invertebrate epifauna was greatest in mature forest, intermediate in 15 yr old forest, and lowest in recently cleared forest, but the reverse was the case for invertebrate in-fauna, with highest densities in recently cleared sites (Sasekumar and Chong, 1998). These results are supported by an investigation of the fishes of a restored mangrove area in Florida where fish diversity among the oldest mangrove habitats was signifi-cantly higher than among more recently planted habitats (Barimo and Serafy, 2003). These patterns likely significantly influence the distribution patterns and species composition of fishes in mangroves, and furthermore, suggest that there may be a succession pattern of fishes colonizing mangroves.

The question of scale is important when discussing diversity of mangrove fishes and their interrelationships with adjacent habitats. The very large areas of estua-rine waters and their associated mangroves extending over 1000s of km2 of sea in Southeast Asia, West Africa, and Northeast South America contrast markedly with the smaller mangrove systems of estuaries along the East African, East Australian, and Caribbean coasts. The marked contrast between clear reef waters and turbid mangrove waters so characteristic of the latter are not evident in the Indo-West Pa-cific where the fishes live in a more uniformly turbid and low salinity environment (Blaber, 2000) and where linkages between mangrove structure, depth, and current speed may be more influential. On a smaller scale, further indication of the impor-tance of linkages has been demonstrated recently for fishes in sub-tropical Moreton Bay in Queensland using a novel hierarchical landscape approach (Pittman et al., 2004). They quantified substratum structure at three spatial scales: whole landscape mosaic (10s of ha); habitat type (100s m2 to 1 ha); and within patch scale (cm2 to m2) and using fish densities, showed that more species used mangroves with adjacent continuous seagrass beds than used mangroves with adjacent patchy seagrass or


unvegetated mudflats. Highest assemblage densities were within spatially heteroge-neous mangroves, but larger bodied and schooling species (e.g., Sillago spp.) exhib-ited a preference for fragmented mangroves with open water areas with high edge, high light penetration, and low shoot density that allowed unimpeded swimming and foraging. In addition, the proportion of mud in surficial sediments was the most important within-patch variable for both density and number of species using man-groves, particularly for gobiids and pleuronectids.

Dependence.—The dependence or otherwise of tropical fishes on mangroves is a question that is not only of importance for understanding ecosystem functioning, but is one that is increasingly being asked by fisheries and environmental managers as fish catches decline and the worldwide area of mangroves decreases. Estuarine dependence of fishes has been extensively reviewed and discussed and detailed infor-mation is available, for example, in Lenanton and Potter (1987), Day et al. (1989), Pot-ter et al. (1990), Whitfield (1994, 1998), Blaber (1997, 2000), and Nordlie (2003). The estuarine dependence paradigm was first developed for fishes of the southeastern USA and southern Africa, primarily from studies of temperate and warm temperate estuaries. In these non-tropical, mainly non-mangrove areas, many species have been shown to be dependent on the estuarine environment during their juvenile phase (Day et al., 1981; Deegan and Thompson, 1985). Unfortunately, the paradigm gained too ready acceptance by workers everywhere, and in the case of tropical mangrove-fish associations, often without sufficient critical evidence. Longhurst and Pauly (1987) questioned the degree of estuarine dependence of sub-tropical and tropical coastal fishes and concluded that out of 20 areas examined, estuarine dependence be demonstrated in only two areas: in South Africa and southern Brazil, both in non-mangrove systems. Blaber and Blaber (1980) and Blaber (1981) postulated that most of the estuarine fishes of the tropics are not estuarine per se, but are character-istic of shallow turbid areas, often with variable salinity. These areas occur over very large areas of the sea in South and Southeast Asia, West Africa, and northern South America as well as in estuaries, but are restricted to estuaries in East Africa, eastern Australia, and parts of the USA. All of these areas and their fishes coincide with regions where mangroves are a primary habitat type. Variations in the complexity of mangrove habitats and the high species diversity of fishes makes any generaliza-tions about dependency difficult. Nevertheless, many species are dependent on the mangrove environment for all or part of their life cycle. Three main hypotheses have been advanced to explain such dependency: (1) reduced predation linked to depth, structure, and turbidity; (2) increased food supply for post-larvae and juveniles; and (3) shelter in quiet (non-turbulent) waters for post-larvae and juveniles. Numbers of fish species that are dependent on the mangrove environment varies both taxo-nomically and zoogeographically. For example, in most tropical regions, juveniles of most species of Mugilidae are found mainly in sheltered estuarine/mangrove wa-ters, whereas among the Carangidae, juveniles of only some species are estuarine-dependent and most are not. Comparing regions, about 30% of the species in the Embley estuary in northern Australia and a similar percentage in the Lupar estuary in Sarawak, Malaysia, are estuarine/mangrove dependent (Blaber et al., 1989). Al-though similar percentages of estuarine-dependent species are also recorded in the tropical east Atlantic in West African mangrove areas (Baran, 1995), the situation in the tropical west Atlantic appears to be different. In the Caete estuary in northeast Brazil, at least 70% of species are estuarine-dependent (Barletta et al., 2003, 2005).


It is important, to note however, that some tropical species that are estuarine-de-pendent, are apparently not mangrove-dependent. A good example here would be the economically important tropical shads (genus Tenualosa) of south and southeast Asia (Blaber et al., 2003).

There has been much discussion about the role of mangrove habitats as nurseries or feeding grounds for fish from adjacent habitats. As with species composition, the role of mangroves in this regard is apparently highly variable. In a series of studies in the Caribbean it has been shown unequivocally that many reef fish species utilize mangroves as nurseries. These studies are particularly interesting because they refer mainly to non-estuarine mangroves and give some insight into whether fishes also depend on mangroves in non-estuarine conditions. These non-estuarine mangroves can have higher densities of juvenile fishes than adjacent seagrass beds, channels, and mudflats, but similar densities to those on coral reefs (Nagelkerken and van der Vel-de, 2002). Different patterns of abundance of juveniles in the mangroves are thought to be related to the degree of structural complexity. The juveniles of at least 17 spe-cies of Caribbean reef fish are highly associated with bays containing mangroves and seagrass beds and are absent in bays without these habitats (Nagelkerken et al., 2000, 2002, 2004a; Dorenbosch et al., 2004). The situation is not straightforward, how-ever, and interestingly, Nagelkerken et al. (2004b) demonstrated using stable carbon and nitrogen isotope analyses that mangroves in the Caribbean are not important feeding grounds for juvenile reef fish from adjacent seagrass beds. They analyzed 23 species of which only five showed signatures indicative of food intake from the mangroves. The patterns of habitat use in five habitats in a marine embayment in Zanzibar showed a high similarity to those in the Caribbean (Lugendo et al., 2005). However, caution is needed in equating any lack of mangrove isotope signals in fishesequating any lack of mangrove isotope signals in fishes to a lack of food utilization from mangrove habitats. Epiphytes growing on man-groves or their pneumatophores may be an important food source, both directly and indirectly, for fishes entering mangrove environments and this carbon would have a different signal from mangrove carbon. Results from Biscayne Bay, Florida lend onlyResults from Biscayne Bay, Florida lend only partial support to an obligatory ontogenetic movement between reef and mangrove, with only two of five species examined conforming to this paradigm (Serafy et al., 2003). However, working in Belize and Mexico, Mumby et al. (2004) showed that mangroves were unexpectedly important as intermediate nursery habitats for reef fish and strongly influenced the community structure of fish on neighboring coral reefs. They showed that the biomass of four commercially important species more than doubled when the adult habitat was connected to mangroves, and that one spe-cies, Scarus guacamaia Cuvier, 1829 is functionally dependent on mangroves as a juvenile and has suffered local extinctions after mangrove removal. Recent experi-mental results from the Caribbean emphasize the different responses by juveniles of various reef species to the structure, food, and shade offered by mangroves, but clarify that the presence of these features contributes significantly to their attractive-ness to juvenile reef fish (Verweij et al., 2006).

In the large estuarine areas of south and southeast Asia, extensive mangrove for-ests are not often in close proximity to coral reefs, as they are in the Caribbean, so most ontogenetic habitat shifts relate to movements between coastal ocean and estuary or shallow to deeper water (Blaber, 1997). There are exceptions, however: Chong et al. (2005) found that the high diversity of fishes in the mangroves of Lang-kawi Island in northwest Malaysia is strongly influenced by the presence of coral


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reefs, limestone karst reefs, and seagrass meadows adjacent to the mangroves, that the fish fauna contains many species not usually found in Indo-Pacific mangroves, and that there is considerable movement among the habitats; a situation that mirrors that of the Caribbean. In the South Pacific, however, where relatively small areas of mangroves often occur adjacent to coral reefs, there appears to be little connectivity between the two habitats although the mangroves may be used as feeding grounds by visiting mobile piscivorous species. Quinn and Kojis (1985) showed that mangroves in northeast Papua New Guinea are nursery areas for a few species of coral reef fish. Similarly, Blaber and Milton (1990) in a study of 13 small Solomon Islands mangrove estuaries showed that these systems had an insignificant role as nurseries for coral reef fishes, Dorenbosch et al. (2005) obtained similar results in east Africa.

Mangrove-Fisheries Connectivity.—It is in the area of fisheries production, including commercial, artisanal, and recreational, that the dogma concerning man-groves and fish has really taken hold. It is now a widely held view by fisheries man-agers, conservationists, and the general public that mangroves are essential for the maintenance of fish populations. There is no doubt that this has had very positive benefits, and given the popularity of recreational fishing in developed countries such as Australia and the USA, has immeasurably helped with the conservation of man-grove environments. However, in order to defend the value of mangroves, such as in debates about alternative land-use, good data are required, and unfortunately, the degree to which many fisheries in the tropics are dependent on mangroves is often unclear.

It is important to distinguish between “fisheries within mangrove systems”, usu-ally of an artisanal or subsistence nature in developing countries, and “offshore (of mangroves) fisheries” that are usually commercial or industrial concerns. In the for-mer case, the activities by traditional or artisanal fishermen may be long-established and are totally dependent on the existence of the mangrove system. Many of these fisheries have been studied in detail and examples are given in Blaber (2000), Jhin-gran (2002), Islam and Haque (2004), and Kathiresan and Qasim (2005). Although the major threats to mangrove fishes are usually linked to environmental degrada-tion, such as removal of mangroves, there is also evidence to suggest that many fish species in tropical estuaries, particularly in south and southeast Asia, are declining in abundance primarily as a result of overfishing. Such overfishing is strongly linked to issues of food security, but unlike many other human activities in and around tropical estuaries, fisheries are almost completely dependent on the maintenance of ecosystem integrity (Blaber, 2005).

Links between offshore fisheries production and mangrove loss are much harder to quantify, if only because the large scale removal of mangroves for aquaculture and development purposes has coincided with increased fishing pressure and more efficient fishing technologies.

Much of the evidence about the importance of mangroves to fisheries production has come from studies of penaeid prawns. Research in the Gulf of Mexico (Turner, 1977), Indonesia (Martosubroto and Naamin, 1977), India (Kathiresan and Rajen-dran (2002), Australia (Staples et al., 1985; Vance et al., 1996), and the Philippines (Primavera, 1998), provides good evidence that there is a correlation between com-mercial offshore prawn catches and the total area of adjacent mangroves. Pauly and Ingles (1986) concluded that most of the variance in the catches of penaeids could be explained by a combination of mangrove area and latitude (Table 2).


The significance of the relationships between fishes and mangroves is rather more equivocal and Robertson and Blaber (1992) concluded that in spite of the correla-tions between mangrove area and commercial fish catches, a causal link has not been established experimentally. In the Gulf of Mexico, Yanez-Arancibia et al. (1985) showed a positive correlation between commercial fish catches and mangrove area, and Barbier and Strand (1998) determined the effects of changes in mangrove area in the Laguna de Terminos on the production and value of prawn harvests in Campeche from 1980 to 1990. In Vietnam, De Graaf and Xuan (1998) demonstrated a similar relationship and indicated that 1 ha of mangrove forest supports a marine catch of 450 kg yr–1. The few studies that have quantified relationships between mangroves and coastal resources were summarized by Baran (1999) and Manson et al. (2005), but it must be reiterated that these are correlations and that causal links have not been established experimentally (see Table 2). In recent reviews of the subject, Baran (1999) and Baran and Hambrey (1999), showed that all of 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 avail-ability, contributing to the relationships. These reviews also show that finding a rela-finding a rela-tionship between mangrove area and fish production is not straightforward, because (i) closely related fish species can have very different ecological requirements, which blurs possible global relationships; (ii) results drawn from site-specific studies can-not be generalized to large areas in different geomorphic and climatic settings; and (iii) fishery statistics, in most cases, cannot be disaggregated enough to link catches to specific mangrove zones. More recently, in a review of the role of mangroves asMore recently, in a review of the role of mangroves as nursery habitats for transient fishes and decapods, Sheridan and Hays (2003) con-cluded that the case for identifying flooded mangrove forests as critical nursery habi-tat remains equivocal until sufficient further experimental and quantitative studies have been carried out. Furthermore, in Queensland, Australia, Manson et al. (2005) demonstrated an empirical link between the extent of mangrove habitat and fishery production (mainly Crustacea) for three mangrove-related species, but such links for four non-mangrove estuarine species were less significant, and for these species, latitude was the dominant variable.

As indicated earlier, not all mangroves or areas in mangrove systems, have the same relationships with fishes. For example, Vance et al., 1996 demonstrated that in northern Australia, the mangrove forests fringing deeper water contain much of the functionality compared with the more inland shallower, intertidal areas. There are other studies that showed that the inland shallower area of the mangrove forests are preferred areas by shrimps and small fishes (Rönnbäck et al., 1999; Affendy and Chong, 2007) presumably to avoid predation. 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. Kapetsky (1985) suggested that much of the functional value of mangroves might be retained from a smaller area of man-groves, for example, 75% of nursery function from 50% of original area. However, it is the deeper fringing areas of mangroves adjacent to the ocean that have been most attractive and suitable for aquaculture pond development throughout the tropics and have hence suffered greater proportional losses than more inland mangroves. This is not to say that shallower inland mangroves may not be important, particularly with regard to small fish species diversity and conservation, as demonstrated by Taylor et al. (1995) in Florida and Rönnbäck et al. (1999) in the Philippines. Loneragan et al.


(2005) suggested that landings of penaeids in west peninsular Malaysia have been maintained or increased despite large losses of mangroves. However, a recent review by Chong (2006) indicated that landings of penaeids on the west coast of peninsular Malaysia had actually declined from 60,967 t in 1989 to 39,296 t in 2003 (35% reduc-tion) despite a reduction in fishing effort, as their main nursery habitat (mangroves) shrunk in area (23% loss). In the tropical Atlantic, recent quantitative data from the Caribbean suggest that the current rates of mangrove deforestation will have serious consequences for coral reef fish and fisheries (Mumby et al., 2004).

Costanza et al. (1997) calculated that the economic value of estuaries in terms of services and natural capital per hectare was the highest of all ecosystems. Tropical mangrove systems, in particular, are zones of high productivity (Blaber, 2000), and assuming that most of the fisheries productivity is closely linked to mangroves, a number of recent studies have emphasized the economic value of mangroves, espe-cially in the developing world (Hamilton et al., 1989; Barbier and Strand, 1998; Nick-erson, 1999; Barbier, 2000). Rönnbäck (2001) stated that “one major driving force behind the loss of more than 50% of the world’s mangroves during the last decade is the inability among economists to recognize and value all goods and services pro-duced by this ecosystem.” In estimating the welfare effects of mangrove-fisheries linkages in Thailand, Barbier et al. (2002) commented that the fisheries most likely to be affected by habitat losses and impacts are those containing a high proportion of artisanal fishers.

Evidence of the value of mangroves to fish and fisheries is rapidly increasing and appears almost overwhelming, but the case at the moment is not proven because much of the evidence is circumstantial. Therefore, despite the fact that the relation-ship has gained widespread scientific and public acceptance, there is still an urgent need for experimental and quantitative studies [such as that of Verweij et al. (2006)] to support the dogma, and to lend weight to the economic arguments that the value of retaining mangroves far exceeds the value of their destruction for whatever pur-pose. Mangroves are still being lost at an unacceptable rate from all points of view—ecological, economic, conservation, and human safety (e.g., Tsunami protection).

Acknowledgments

I am very grateful to the organizers of the “First International Symposium on MangrovesFirst International Symposium on Mangroves as Fish Habitat” at the Rosenthiel School of Marine and Atmospheric Science, University of Miami, for providing the stimulus for writing this paper and the funds for presenting it at the conference.

Literature Cited

Affendy, N. and V. C. Chong. 2007. Shrimp ingress into mangrove forests of different age stands, Matang Mangrove Forest Reserve, Malaysia. Bull. Mar. Sci. 80: 915.

Albaret, J.J. 1994. Les poissons: biologie et peuplements. Pages 239–279Pages 239–279 in J. R. Durand, Du-four, D. Guiral and Zabi, eds. Environnement et ressources aquatiques de C�te d’Ivoire,Environnement et ressources aquatiques de C�te d’Ivoire, tome II - les mileux lagunaire. Editions de L’ORSTOM, Paris.

Baran, E. 1995. Dynamique spatio-temporelle des peuplements de poissons estuariens en Gui-née - relations avec les milieu abiotique. PhD Thesis, Université de Bretagne Occidentale, France.


_______. 1999. A review of quantified relationships between mangroves and coastal resources.and coastal resources. Phuket Mar. Biol. Centre, Res. Bull. 62: 57–64.

_______. 2000. Biodiversity of estuarine fish faunas in West Africa. Naga, The ICLARM Quar-terly 23: 4–9.

_______ and J. Hambrey. 1999. Mangrove conservation and coastal management in Southeast Asia: What impact on fisheries resources. Mar. Poll. Bull. 37 : 431–440.Mar. Poll. Bull. 37 : 431–440.

Barbier, E. B. 2000. Valuing the environment as input: review of applications to mangrove fish-ery linkages. Ecol. Econ. 35: 47–61.

__________ and I. Strand. 1998. Valuing mangrove-fishery linkages. Environ. Resour. Econ. 12: 151–166.

__________, _________, and S. Sathirathai. 2002. Do open access conditions affect the valua-tion of an externality? Estimating the welfare effects of mangrove-fishery linkages in Thai-land. Environ. Resour. Econ. 21: 343–367.

Barimo, J. F. and J. E. Serafy. 2003. Fishes of a restored mangrove habitat on Key Biscayne, Florida. Fla. Sci. 66: 12–22.

Barletta, M., A. Barletta-Bergen, U. Saint-Paul, and G. Hubold. 2003. Seasonal changes in den-sity, biomass, and diversity of estuarine fishes in tidal mangrove creeks of the lower Caeté Estuary (northern Brazilian coast, east Amazon). Mar. Ecol. Prog. Ser. 256: 217–228.

__________, __________, __________, and _________. 2005. The role of salinity in structuring the fish assemblages in a tropical estuary. J. Fish Biol. 66: 45–72.

Blaber, S. J. M. 1980. Fishes of the Trinity Inlet system of North Queensland with notes on the ecology of fish faunas of tropical Indo-Pacific estuaries. Aus. J. Mar. Freshw. Res. 31: 137–146.

___________. 1981. The zoogeographical affinities of estuarine fishes in southeast Africa. S. Afr. J. Sci. 77: 305–307.

___________. 1997. Fish and Fisheries of Tropical Estuaries. Chapman and Hall, London. 367 p.___________. 2000. Tropical Estuarine Fishes: Ecology, Exploitation and Conservation, Black-

well, Oxford, 372 p. __________. 2007. Fisheries management and conservation in tropical estuaries–is a balance

possible? Proc. 4th Word Fisheries Congress, Vancouver.__________ and T. G. Blaber. 1980. Factors affecting the distribution of juvenile estuarine and

inshore fish. J. Fish Biol. 17: 143–162.__________ and D. P. Cyrus. 1981. A revised checklist and further notes on the fishes of the

Kosi system. Lammergeyer 31: 5–14.__________ and D. A. Milton. 1990. Species composition, community structure and zoogeog-

raphy of fishes of mangroves in the Solomon Islands. Mar. Biol. 105: 259–268.___________, D. T. Brewer, and J. P. Salini. 1989. Species composition and biomasses of fishes in

different habitats of a tropical northern Australian estuary: their occurrence in the adjoin-ing sea and estuarine dependence. Est. Coast. Shelf Sci. 29: 509–531.

___________, __________, and ________. 1994. Comparisons of the fish communities of tropi-cal estuarine and inshore habitats in the Gulf of Carpentaria, northern Australia. Pages 363–372 in K. R. Dyer and R. J. Orth, eds. Int. Symp. Series, ECSA 22, Olsen and Olsen, Denmark.

__________, D. A. Milton, D. T. Brewer, and J. P. Salini. 2003. Biology, fisheries and status of tropical shads (Tenualosa) in south and southeast Asia. Am. Fish. Soc. Symp. 35: 49–58.

Boeseman, M. 1963. An annotated list of fishes from the Niger Delta. Zool. Verh. 61: 1–54.Boujard, T. and R. Rojas-Beltran. 1988. Longitudinal zonation of the fish population of the Sin-

namary River (French Guiana). Revue d’hydrobiologie tropicale 21: 47–61.Revue d’hydrobiologie tropicale 21: 47–61.Cervigón, F. 1985. The ichthyofauna of the Orinoco estuarine water delta in the west Atlan-

tic coast, Caribbean. Pages 57–78 in A. Yáñez-Arancibia, ed. Fish Community Ecology in Estuaries and Coastal Lagoons: Towards an Ecosystem Integration. UNAM Press, MexicoUNAM Press, Mexico City.


Chaves, P. and J. L. Bouchereau. 1999. Biodiversité et dynamique des peuplements ichtyiques de la mangrove de Guaratuba, Bresil. Oceanol. Acta 22: 353–364.

Chong, V. C. 2006. Importance of coastal habitats in sustaining the fisheries industry. PagesImportance of coastal habitats in sustaining the fisheries industry. Pages 1–28 in National Fisheries Symp. on “Advancing R and D Towards Fisheries Business Op-portunities”, 26–28 June 2006, Crown Plaza Riverside Hotel, Kuching, Sarawak, Malaysia.

__________, A. Sasekumar, M. U. C. Leh, and R. D’Cruz. 1990. The fish and prawn communi-1990. The fish and prawn communi-ties of a Malaysian coastal mangrove system, with comparisons to adjacent mud flats andMalaysian coastal mangrove system, with comparisons to adjacent mud flats and inshore waters. Est. Coast. Shelf Sci. 31: 703–722.

__________, N. Affendy, A. L. Ooi, and L. L. Chew. 2005. The fishes and invertebrates of man-grove estuaries in northeastern Langkawi Island. Malay. Nat. J. 57: 193–208.

Chua, Thia-Eng. 1973. An ecological study of the Ponggol estuary in Singapore. Hydrobiologia 43: 505–533.

Costanza, R., R. d’Arge, R. de Groot, S. Farber, M. Grasso, B. Hannon, K. Limburg, S. Naeem, R. V. O’Neill, J. Paruelo, R. G. Raskin, P. Sutton, and M. van den Belt. 1997. The value of the world’s ecosystem services and natural capital. Nature 387: 253–260.

Day, J. H. 1974. The ecology of Morrumbene estuary, Moçambique. Trans. Roy. Soc. S. Afr. 41: 43–97.

_______, S. J. M. Blaber, and J. H. Wallace. 1981. Estuarine Fishes. Pages 97–221 in J. H. Day, ed. Estuarine Ecology with particular reference to southern Africa A. A. Balkema, Cape Town, South Africa.

Day, J. W., Jr., C. A. S. Hall, W. M. Kemp, and A. Yáñez-Arancibia. 1989. Estuarine Ecology. John Wiley and Sons, New York. 576 p.

De Graaf, G. J. and T. T. Xuan. 1998. Extensive shrimp farming, mangrove clearance and marineExtensive shrimp farming, mangrove clearance and marine fisheries in the southern provinces of Vietnam. Mangroves and Saltmarshes 2: 159–166.

Deegan, L. A. and B. A. Thompson. 1985. The ecology of fish communities in the Mississippi River deltaic plain. Pages 35–56 in A. Yáñez-Arancibia, ed. Fish Community Ecology in Estuaries and Coastal Lagoons: Towards an Ecosystem Integration. UNAM Press, Mexico City.

Dorenbosch, M., M. C. van Riel, I. Nagelkerken, and G. van der Velde. 2004. The relationship2004. The relationship of fish densities to the proximity of mangrove and fish nurseries. Est. Coast. Shelf Sci. 60: 37–48.

____________, M. G. G. Grol, J. A. Christianen, I. Nagelkerken, and G. van der Velde. 2005. Indo-Pacific seagrass beds and mangroves contribute to fish density and diversity on adja-cent coral reefs. Mar. Ecol. Prog. Ser. 302: 63–76.

Fagade, S. O. and C. I. O. Olaniyan. 1972. The biology of the West African shad, Ethmalosa fimbriata (Bowdich) in the Lagos Lagoon. J. Fish Biol. 4: 519–533.

FAO. 2003. Status and trends in mangrove area extent worldwide. By Wilkie, M. L. and For-tuna, S. Forest Resources Assessment Working Paper No. 63. Forest Resources Division. FAO, Rome. (Unpublished)

Gilbert, C. R. and D. P. Kelso. 1971. Fishes of the Tortuguero area, Carribbean Costa Rica. Bull.Bull. Fl State Mus. Biol. Sci. 16: 1–54.

Haines, A. K. 1979. An ecological survey of fish of the lower Purari River system, Papua New Guinea. Department of Minerals and Energy, Papua New Guinea, Purari River Hydroelec-tric Scheme Environmental Studies 6: 1–102.

Hamilton, L., J. Dixon, and G. Miller. 1989. Mangroves: an undervalued resource of the land and sea. Ocean Yearb. 8: 254–288.

Harrison, T. D. and A. K. Whitfield. 1995. Fish community structure in three temporarily open/closed estuaries on the natal coast. Ichthyol. Bull. J. L. B. Smith Inst. Ichthyol. 64: 1–80.

Hayase, S. and M. Fadzil. 1999. Fish distribution and abundance in Matang and Merbok man-grove waters, west coast of peninsula Malaysia: II. Preliminary results for 1996–1997. Pages 59–91 in V. C. Chong and P. S. Choo, eds. Productivity and sustainable utilization of brack-ish water mangrove ecosystems. Japan International Research Center for Agricultural Sci-ences, Tokyo, Japan.


Hutchings, P. and P. Saenger. 1987. Ecology of Mangroves, University of Queensland Press, St Lucia, Queensland, Australia.

Inger, R. E. 1955. Ecological notes on the fish fauna of a coastal drainage of North Borneo. Fieldiana; Zoology 37: 47–90.

Islam, M. S. and M. Haque. 2004. The mangrove-based coastal and nearshore fisheries of Ban-gladesh: ecology, exploitation and management. Rev. Fish Biol. Fish. 14: 153–180.

Jhingran, V. G. 2002. Fish and fisheries of India (3rd ed). Hindustan Publishing Corporation, New Delhi. 728 p.

Jones, S. and K. H. Sujansingani. 1954. The fish and fisheries of the Chilka Lake with statistics of fish catches for the years 1948–1950. Ind. J. Fish. 1: 256–344.

Kapetsky, J. M. 1985. Mangroves, fisheries and aquaculture. FAO Fisheries Report 338: 17–36.Kathiresan, K. and N. Rajendran. 2002. Fishery resources and economic gain in three mangrove

areas on the south-east coast of India. Fish. Manag. Ecol. 9: 277–283.___________ and S. Z. Qasim. 2005.Biodiversity of Mangrove Ecosystems. Hindustan Publish-

ing Corporation, New Delhi. Khoo, H. K. 1989. The fisheries in the Matang and Merbok mangrove ecosystems. Pages 147–

169 in S. M. Phang, A. Sasekumar, and S. Vickineswary, eds. Proc. 12th Annual Seminar of the Malaysian Society of Marine Sciences, Kuala Lumpur, Malaysia.

Krishnamurthy, K. and M. J. Jeyaseelan. 1981. Early life history of fishes from Pichavaram man-1981. Early life history of fishes from Pichavaram man-grove ecosystem of India. Rapports et procés verbaux des Réunions, Conseil PermanentRapports et procés verbaux des Réunions, Conseil Permanent International pour l’exploration de la Mer 178: 416–423.

Lenanton, R. C. J. and I. C. Potter. 1987. Contribution of estuaries to commercial fisheries, R. C. J. and I. C. Potter. 1987. Contribution of estuaries to commercial fisheries in temperate Western Australia and the concept of estuarine dependance. Estuaries 10: 28–35.

León, R. A. and J. B. Racedo. 1985. Composition of fish communities in the lagoon and estua-rine complex of Cartagena Bay, Ciénaga de Tesca and Ciénaga Grande de Santa Marta, Co-lombian Caribbean. Pages 536–556 in A. Yáñez-Arancibia, ed. Fish Community Ecology in Estuaries and Coastal Lagoons: Towards an Ecosystem Integration. UNAM Press, Mexico.

Lin, H. J. and K. T. Shao. 1999. Seasonal and diel changes in a subtropical mangrove fish as-semblage. Bull. Mar. Sci. 65: 775–794.

Little, M. C., P. J. Reay, and S. J. Grove. 1988. Distribution gradients of ichthyoplankton in an East African Mangrove creek. Est. Coast. Shelf Sci. 26: 669–677.

Loneragan, N. R., N. Ahmad Adnan, R. M. Connolly, and F. J. Manson. 2005. Prawn landings and their relationship with the extent of mangroves and shallow waters in western penin-sula Malaysia. Est. Coast. Shelf Sci. 63: 187–200.

Longhurst, A. R. and D. Pauly. 1987. Ecology of Tropical Oceans, Academic Press, San Diego.Lugendo, B. R., A. Pronker, I. Cornelissen, A. de Groene, I. Nagelkerken, M. Dorenbosch, G. vanrnelissen, A. de Groene, I. Nagelkerken, M. Dorenbosch, G. van

der Velde, and Y. D. Mgaya. 2005. Habitat utilisation by juveniles of commercially impor-tant fish species in a marine embayment in Zanzibar, Tanzania. Aquat. Living Resour. 18: 149–158.

Lyons, J. and D. W. Schneider. 1990. Factors influencing fish distribution and community struc-ture in a small coastal river in southwestern Costa Rica. Hydrobiologia 203: 1–14.

Manson, F. J., N. R. Loneragan, B. D. Harch, G. A. Skilleter, and L. Williams. 2005. A broad-scale analysis of links between coastal fisheries production and mangrove extent: A case study for northeastern Australia. Fish. Res. 74: 69–86.

Martosubroto, P. and N. Naamin. 1977. Relationship between tidal forests (mangroves) and commercial shrimp production in Indonesia. Mar. Res. Indones. 18: 81–86.

Mumby, P. J., A. J. Edwards, J. E. Arias-González, K. C. Lindeman, P. G. Blackwell, A. Gall, M. I. Gorczynska, A. R. Harborne, C. L. Pescod, H. Renken, C. C. C. Wabnitz, and G. Llewellyn. 2004. Mangroves enhance the biomass of coral reef fish communities in the Caribbean. Nature 427: 533–536.


Nagelkerken, I. and G. van der Velde. 2002. Do non-estuarine mangroves harbour higher densi-ties of juvenile fish than adjacent shallow-water and coral reef habitats in Curacao (Nether-lands Antilles)? Mar. Ecol. Prog. Ser. 245: 191–204.

____________ and _____________. 2004a. Relative importance of interlinked mangroves and seagrass beds as feeding habitats for juvenile reef fish on a Caribbean island. Mar. Ecol. Prog. Ser. 274: 153–159.

____________ and _____________. 2004b. Are Caribbean mangroves important feeding grounds for juvenile reef fish from adjacent seagrass beds. Mar. Ecol. Prog. Ser. 274: 143–151.

____________, C. M. Roberts, G. van der Velde, M. Dorenbosch, M. C. van Riel, E. Cocheret de la Morinière, and P. H. Nienhuis. 2002. Mar. Ecol. Prog. Ser. 244: 299–305.

____________, G. van der Velde, M. W. Gorissen, G. J. Meijer, T. van’t Hof, and C. den Har-tog. 2000. Importance of mangroves, seagrass beds and the shallow coral ref. as a nursery for important coral reef fishes, using a visual census technique. Est. Coast. Shelf Sci. 51: 31–44.

Nickerson, D. J. 1999. Trade-offs of mangrove area development in the Philippines. Ecol. Econ. 28: 279–298.

Nordlie, F. G. 2003. Fish communities of estuarine saltmarshes of eastern North America, and comparisons with temperate estuaries of other continents. Rev. Fish Biol. Fish. 13: 281–325.

Paranagua, M. N. and E. Eskinazi-Leça. 1985. Ecology of a northern tropical estuary in Brazil1985. Ecology of a northern tropical estuary in Brazil and technological perspectives in fish culture. Pages 595–614 in A. Yáñez-Arancibia, ed. Fish Community Ecology in Estuaries and Coastal Lagoons: Towards an Ecosystem Inte-gration. UNAM Press, Mexico.

Pauly, D. and J. Ingles. 1986. The relationship between shrimp yields and intertidal vegetation (mangrove) areas: a reassessment. Pages 227–284 in A. Yáñez-Arancibia and D. Pauly, eds. IOC/FAO Workshop on Recruitment in Tropical Coastal Demersal Communities. Ciudad de Carmen, Mexico. IOC, UNESCO, Paris.

Pinto, L. 1988. Population dynamics and community structure of fish in the mangroves of Pag-bilao, Philippines. J. Fish Biol. 33: 35–43.

Pittman, S. J., C. A. McAlpine, and K. M. Pittman. 2004. Linking fish and prawns to their envi-ronment: a hierarchical landscape approach. Mar. Ecol. Prog. Ser. 283: 233–254.

Potter, I. C., L. E. Beckley, A. K. Whitfield, and R. C. J. Lenanton. 1990. Comparisons between the roles played by estuaries in the life cycles of fishes in temperate western Australia and southern Africa. Environ. Biol. Fish. 28: 143–178.

Primavera, J. H. 1998. Mangroves as nurseries: shrimp populations in mangrove and non-man-grove habitats. Est. Coast. Shelf Sci. 46: 457–464.

Quinn, N. J. and B. L. Kojis. 1987. The influence of diel cycle, tidal direction and trawl align-ment on beam trawl catches in an equatorial estuary. Environ. Biol. Fish. 19: 297–308.

Robertson, A. I. and S. J. M. Blaber. 1992. Plankton, epibenthos and fish communities. Pages 173–224 in A. I. Robertson and D. M. Alongi, eds. Tropical Mangrove Ecosystems. Ameri-can Geophysical Union, Washington DC, USA.

____________ and N. C. Duke. 1990. Mangrove fish communities in tropical Australia: spa-tial and temporal patterns in densities, biomass and community structure. Mar. Biol. 104: 369–379.

Rönnbäck, P. 2001. Ecological economics of fisheries supported by mangrove ecosystems: eco-nomic efficiency, mangrove dependence, sustainability and benefit transfer. 2nd Western Indian Ocean Science Association Scientific Symp.–Book of Abstracts. 41 p.

___________, M. Troell, N. Kautsky, and J. H. Primavera. 1999. Distribution pattern of shrimps and fish among Avicennia and Rhizophora microhabitats in the Pagbilao mangroves, Philip-pines. Est. Coast. Shelf Sci. 48: 223–234.

Sasekumar, A. and V. C. Chong. 1998. Faunal diversity in Malaysian mangroves. Global Ecol. Biogeogr. Lett. 7: 57–60.


____________ and C. R. Wilkinson. 1994. Compatible and incompatible uses of mangroves in ASEAN. Pages 77–83 in C. R. Wilkinson, ed. Proc. Third ASEAN-Australia Symp. on Liv-ing Coastal Resources, Bangkok, Thailand. Australian Institute of Marine Science, Towns-ville, Australia.

___________, V. C. Chong, K. H. Lim, and H. Singh. 1994. The fish community of Matang waters. Pages 457–464 in S. Sudara, C. R. Wilkinson, and L. M. Chou, eds. Proc. Third ASEAN-Australia Symp. on Coastal Living Resources, vol. 1: Status reviews. Chulalong-korn University, Bangkok, Thailand.

Serafy, J. E., C. H. Faunce, and J. J. Lorenz. 2003. Mangrove shoreline fishes of Biscayne Bay, Florida. Bull. Mar. Sci. 72: 161–180.

Sheridan, P. and C. Hays. 2003. Are mangroves nursery habitat for transient fishes and deca-pods. Wetlands 23: 449–458.

Staples, D. J., D. J. Vance, and D. S. Heales. 1985. Habitat requirements of juvenile penaeid prawns and their relationship to offshore fisheries. Pages 47–54 in P. C. Rothlisberg, B. J. Hill, and D. J. Staples, eds. Second Australian National Prawn Seminar, Kooralbyn, Aus-tralia. CSIRO, Cleveland, Queensland, Australia.

Taylor, D. S., W. P. Davis, and B. J. Turner. 1995. Rivulus marmoratus: ecology of distributional patterns in Florida and the central Indian River Lagoon. Bull. Mar. Sci. 57: 202–207.

Thayer, G. W., D. R. Colby, and W. F. Hettler, Jr. 1987. Utilization of the red mangrove prop root habitiat by fishes in south Florida. Mar. Ecol. Prog. Ser. 35: 25–38.

Turner, R. E. 1977. Intertidal vegetation and commercial yields of penaeid shrimps. Trans. Am. Fish. Soc. 106: 411–416.

Vance, D. J., M. D. E. Haywood, D. S. Heales, R. A. Kenyon, N. R. Loneragan, and R. C. Pendrey. 1996. How far do prawns and fish move into mangroves? Distribution of juvenile banana prawns Penaeus merguiensis and fish in a tropical mangrove forest in northern Australia. Mar. Ecol. Prog. Ser. 131: 115–124.

Verweij, M. C., I. Nagelkerken, D. De Graaf, M. Peeters, E. J. Bakker, and G. van der Velde. 2006.2006. Structure, food and shade attract juvenile coral reef fish to mangrove and seagrass habitats: a field experiment. Mar. Ecol. Prog. Ser. 306: 257–268.Mar. Ecol. Prog. Ser. 306: 257–268.

Whitfield, A. K. 1994. Fish species diversity in southern African estuarine systems: an evolu-tionary perspective. Environ. Biol. Fish. 40: 37–48.

____________. 1998. Biology and Ecology of Fishes in Southern African Estuaries. Ichthyol. Bull. J. L. B. Smith Inst. Ichthyol., Grahamstown, South Africa. 223 p.

Yáñez-Arancibia, A. 1978. Taxonomy, ecology and structure of fish communities in coastal lagoons with ephemeral inlets on the Pacific coast of Mexico. Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autónoma de México, Publicaciones Especiales 2: 1–306.

_______________, A. L. Lara-Dominguez, P. Sanchez-Gil, I. Vargas Maldonado, M. De La C. García Abad, H. Álvarez-Guillén, M. Tapia García, D. Flores Hernández, and F. Amezcua Linares. 1985. Ecology and evaluation of fish community in coastal ecosystems: estuary-1985. Ecology and evaluation of fish community in coastal ecosystems: estuary-shelf interrelationships in the southern Gulf of Mexico. Pages 475–498 in A. Yáñez-Aran-cibia, ed. Fish Community Ecology in Estuaries and Coastal Lagoons: Towards an Ecosys-tem Integration. UNAM Press, Mexico City.

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