SciELO - Scientific Electronic Library Online

vol.52 issue6Effects of plant cover on the macrofauna of Spartina marshes in northern BrazilGills of juvenile fish piaractus mesopotamicus as histological biomarkers for experimental sub-lethal contamination with the Organophosphorus Azodrin®400 author indexsubject indexarticles search
Home Pagealphabetic serial listing  

Services on Demand




Related links


Brazilian Archives of Biology and Technology

Print version ISSN 1516-8913

Braz. arch. biol. technol. vol.52 no.6 Curitiba Nov./Dec. 2009 



Feeding ecology of juvenile dog snapper Lutjanus jocu (Bloch and Shneider, 1801) (Lutjanidae) in intertidal mangrove creeks in Curuçá estuary (Northern Brazil)



Dijane Pantoja MonteiroI; Tommaso GiarrizzoI,II,*; Victoria IsaacI

ILaboratório de Biologia Pesqueira - Manejo dos Recursos Aquáticos; Universidade Federal do Pará; Av. Perimetral 2651; 66040-170; Belém - PA - Brasil
IICenter for Tropical Marine Ecology (ZMT); Fahrenheitst, 6, 28359; Bremen - Germany




The diet and feeding ecology of juvenile dog snapper (Lutjanus jocu) were investigated in 92 specimens collected in four intertidal mangrove creeks of Curuçá estuary, Northern Brazil, between September 2003 and July 2004. No significant differences in total length were found between the sampling months. Feeding intensity was high as indicated by high stomach fullness index and low vacuity index. The most important prey was Penaeidae, followed by Grapsidae and Porcellanidae. The diet of juvenile dog snapper showed clear seasonal differences. Specimens from dry season (September and November) and dry / wet season transition (January), were specialists feeding exclusively on Penaeidae. However, the specimens from wet season (March and May) and wet /dry season transition (July) that consumed mainly Grapsidae, Penaeidae and Porcellanidae were generalist. These seasonal changes in diet could be related to variations in food availability.

Key words: estuary; feeding habits; feeding strategy; juvenile fish; Lutjanus jocu; intertidal mangrove creek


A dieta e a ecologia alimentar de juvenis de Lutjanus jocu foram verificadas em 92 espécimes coletados em quatro canais de maré do estuário do rio Curuçá, Norte do Brasil, entre setembro de 2003 e julho de 2004. O comprimento total dos peixes coletados não apresentou diferenças significativas entre os meses amostrados. A intensidade alimentar foi elevada conforme indicado pelos altos valores do índice de repleção estomacal e os baixos valores do índice de vacuidade. A presa mais importante foi Penaeidae, seguida por Grapsidae e Porcellanidae. A dieta de juvenis de L. jocu apresentou diferenças sazonais evidentes. Os espécimes da estação seca (setembro e novembro) e transição seca/chuvosa (janeiro) foram considerados especialistas alimentando-se exclusivamente de Penaeidae. No entanto, os espécimes da estação chuvosa (março e maio) e da transição chuvosa/seca (julho), que alimentaram-se principalmente de Grapsidae, Penaeidae e Porcellanidae, foram considerados generalistas. Esta mudança sazonal na dieta poderia estar relacionada com a disponibilidade do alimento.




Mangroves are widely considered to be an important habitat for fishes, including juveniles of commercially important species (e.g. Morton, 1990; Laegdsgaard and Johnson, 1995; Giarrizzo and Krumme, 2008, 2009). This habitat is thought to provide excellent shelter against predators due to water turbidity, the structural complexity of these biotopes (Parrish, 1989; Robertson and Blaber, 1992), as well as provide high food availability for the high productivity of the mangroves and the associated epi and benthic fauna (Odum and Heald, 1972; Laegdsgaard and Johnson, 2001).

The Lutjanidae family comprises medium to large sized demersal predacious fishes which inhabiting mangroves and seagrass beds during their juvenile and sub-adult stages and when adults, they migrate to rocky or coral reefs to take up permanent residence there (Druzhinin, 1970; Cervigón, 1993; Cocheret et al., 2003).

The snappers have high market values and are commercially exploited in the tropical and subtropical coastal waters (e.g. Serrano-Pinto and Caraveo-Patiño, 1999; Luckhurst et al., 2000; Sheaves and Molony, 2001; Kiso and Mahyam, 2003; Miranda et al., 2003; Kamukuru and Mgaya, 2004; Rezende and Ferreira, 2004; Rojas et al., 2004).

The dog snapper, Lutjanus jocu, is one of the Lutjanidae species targeted by artisanal, small and medium scale fisheries in the Brazilian coastal waters (Rezende et al., 2003). Its distribution extends along the western Atlantic Ocean, from Massachusetts (USA) to the southern Brazilian coast, including Gulf of Mexico and Caribbean Sea (Froese and Pauly, 2006). In spite of its relative abundance and importance to fisheries, few information on biology and ecology of this species are available. In order to understand the role that L. jocu plays in the trophic food web, the present study examined the feeding ecology of the juvenile dog snapper in the intertidal mangrove creeks of Curuçá estuary. Emphasis was placed on the assessment of season-related feeding habits, as well as feeding strategy.



Study area

Sampling was carried out in four intertidal mangrove creeks located in the estuary of the Curuçá River near the city of Curuçá (0° 10'S, 47° 50'W), Pará, Northern Brazil, approximately 160 km north-east of the state capital Belém (Fig. 1). The estuary is a well preserved environment and has been designated by the Ministry of the Environment as a conservation unit ("Reserva Extrativista"). The climate is hot and humid with mean annual rainfall of 2,526 mm (ANA, 2005; n = 16 years, range: 1,085 – 3,647 mm). Salinity changes according to the season, being low during the rainy season, in the first half of the year and attaining values of marine water during the dry season (Giarrizzo and Krumme, 2007). Tides in the region are characterized by a semi-diurnal pattern with tidal range between 4 and 5 m.



Fish sampling

Bimonthly samples of L. jocu were obtained using a fyke net in four creeks from September 2003 to July 2004 on four consecutive days, yielding a total of 24 samples. The net was composed by two wings (20 m long, 6 m deep, with 20 mm stretch-mesh) and a hoop-net (with 13 mm stretch-mesh). The fyke net was set at the mouth of the creeks at daytime slack high water (HW) in the waxing of the moon (neap tide). During ebb tide, the fish were collected until total drainage of the creek and kept on ice.

Laboratory analysis

Each fish examined was measured to the nearest 0.1 cm of total length (TL) and weighed to the nearest <0.11 g of wet body weight (WT). Gonadal stages were recorded according to Vazzoler (1996) and stomach contents were dissected out. The relative contribution of the number of empty stomachs was recorded (vacuity index). Stomach contents were identified to the lowest possible taxon (Figueiredo and Menezes, 1978; Menezes and Figueiredo, 1980; 1985; Cervigón et al., 1992; Melo, 1996; 1999). Prey items were counted and weighed to the nearest 0.0001 g after removing the surface water by blotting on tissue paper.

Diet analysis

The feeding activity of fish was evaluated by fullness index (%FI):

%FI = (Wcont / WT) × 100

Where Wcont is the weight of stomach contents calculated from the difference between the weights of pre-washed and washed empty stomachs.

For the qualitative and quantitative analysis of dietary composition were used:

- Frequency of occurrence (F): represents the number of stomachs in which a food item was found, expressed as the percentage of total number of non-empty stomachs.

- Percentage numerical abundance (N): considers the number of individuals in each food category expressed as a percentage of the total individuals in all food categories.

- Percentage gravimetric composition (W) represents the total wet weight of a food category expressed as a percentage of the overall weight of stomach contents.

The main food items were identified using the index of relative importance (IRI) of Pinkas et al. (1971), as modified by Hacunda (1981):

IRI = F × (N + W)

This index has been expressed as:

%IRI = (IRI / ΣIRI) × 100

To assess the feeding strategy along the annual cycle studied, the modified Costello (1990) graphical method (Amundsen et al., 1996) was used. In this method, the prey-specific abundance (%Pi) (y – axis) was plotted against the frequency of occurrence (F) (x - axis). The prey-specific abundance (Pi) has been expressed as:

%Pi = (Σ Si /Σ Sti) × 100

Where Si is the number of prey i and Sti is the total number of prey in the stomachs containing prey i.

To assess the bimonthly changes in diet breadth of L. jocu we used the Levins' standardized index

(Bi) (Hurlbert, 1978; Krebs, 1989):

Bi = (1/n – 1) × ( 1/Σ jP2ij -1)

where Bi is the Levins' standardized index for predator i; Pij is proportion of diet of predator i that is made up of prey j, and n is number of prey categories.

This index ranges from 0 to 1; low values indicate the diets dominated by few prey items (specialist predators), high values indicate the generalist diets (Krebs, 1989).

Statistical analysis

The bimonthly variation of TL and %FI were tested using a one-way analysis of variance (ANOVA) (Underwood, 1997). ANOVA assumptions of normality (Kolmogorov-Smirnov test) and homoscedasticity (Bartlett's test) were previously tested. In all the tests, p < 0.05 was used for rejecting the null hypothesis.

The data matrix of bimonthly %IRI values of family diet components were square root transformed (Platell and Potter, 2001) and the similarity was calculated using Bry-Curtis similarity (Marshall and Elliott, 1997). A hierarchical agglomerative cluster analysis (using complete linkage) was performed to describe the similarity in feeding habitat among the months.



Sample characterization

A total of 92 individuals of L. jocu were captured between September 2003 and July 2004. Catch was highest in July (n = 27) and lowest in March (n = 4). The dog snapper ranged in size from 8.4 to 31.0 cm TL, with a mean value (± SD) of 16.5 cm (± 4.5). Total length did not change significantly over the months (ANOVA: F = 1.53, p > 0.05), though mean TL value in March (21.5 ± 2.2 cm) was higher than May value (15.2 ± 5.1 cm).

All the catches were represented only by juveniles.

Feeding activity

Of the 92 stomachs analyzed, 4 (4.3%) were empty. The fullness index (%FI) ranged from 0 to 6% with a mean value (± SD) of 3% (± 1.6%). The values of %FI were not influenced by months sampled (ANOVA, F = 1.31, p > 0.05). However, lowest and highest values occurred in July (2.0 ± 1.7%) and May (3.5 ± 1.8%), respectively.

Overall composition of the diet

A total of 732 prey belonging to 44 taxa were identified, with a average number of prey per stomach of 8.3. Identification to species level was possible in several cases, due to the limited stage of digestion of prey. Frequency of occurrence, percentage of numerical abundance, gravimetric composition and index of relative importance values of different groups, genera and species of prey organisms found in the stomachs are shown in Table 1. Penaeidae and Grapsidae were present in highest abundance (N: 37.8% and 17.8%, respectively) and also had the highest frequency of occurrence (F: 61.4% and 60.2%, respectively) in the dog snapper stomachs. In spite of the numerous Penaeidae ingested, they only represented 15.2% of percentage gravimetric composition. Grapsidae and Porcellanidae dominated in biomass (W: 22.1% and 16.2%, respectively). According to percentage of index of relative importance (%IRI), the most important prey was Penaeidae (36.5%), followed by Grapsidae (27.0%) and Porcellanidae (21.4%).



Temporal variation of the diet and feeding strategy

Cluster analysis carried out on IRI data differentiated two groups at a similarity of 55% (Fig. 2a). The first group (A) was composed of specimens from dry season (September and November) and dry / wet season transition (January), with their diet dominated by Penaeidae (Fig. 2b). The second (group B) contained specimens from the wet season (March and May) and wet / dry season transition (July) that consumed mainly Grapsidae, Penaeidae and Porcellanidae (Fig. 2b).



The interpretation of the diagrams of the modified Costello graphical method (Amundsen et al., 1996) is shown in Figure 3a. The plot of prey-specific abundance (%Pi) and frequency of occurrence (F) of the main components of the diet between September 2003 and July 2004, showed a strong specialization towards Penaeidae to have been eaten by more than half the individuals (F = 61%) and to have high contribution in specific abundance (Pi = 54%) (Fig. 3b). However, some teleost prey (e.g. Sciaenidae, Poecilidae, Haemulidae, Ariidae and Carangidae) presented a low F and a low Pi (lower left quadrant), displaying evidence of a generalist strategy.



The independent analysis of feeding strategy between the temporal groups identified by cluster analysis showed for the group A, a similar tendency to that observed in the total population being Penaeidae the dominant prey component (Pi = 62% e F = 80%) (Fig. 3c). The dog snappers of group B (March, May and July) presented most prey with lower contribution in abundance, indicating a generalized feeding strategy (Fig. 3d). The most representative prey families were Grapsidae (Pi = 31% and F = 83%), Penaeidae (Pi = 42% and F = 55%) and Porcellanidae (Pi = 23% and F =58%).

The Levins' standardized index (Bi) was lower in September (Bi = 0.1) and November (Bi = 0.2) indicating a selective diet and narrow niche width (Fig. 4). A higher value of Bi recorded in March (Bi = 0.7) showed a wide trophic niche. Similar patterns were found in the interpretation of the diagrams of the modified Costello graphical method for each month (no shown).




Population characteristics

The small size of L. jocu specimens caught in intertidal mangrove creeks of Curuçá estuary supported the hypothesis that the mangroves are a nursery ground habitat for juveniles and sub-adults snapper species (Sheaves, 1995; Nagelkerken et al., 2000; Cocheret de la Moriniére et al., 2003). The highest abundances of dog snapper in dry season suggested that the salinity variations and the hydrodynamic regime established by the seasonal changes in freshwater runoff, have been important control factors in the use of mangrove habitat by this species.

Feeding activity

Juvenile dog snapper showed a feeding activity similar to other estuarine fishes (Krumme et al., 2005; Giarrizzo and Saint-Paul, 2008). The high %FI and the limited stage of digestion of prey indicated that the feeding activity started at high tide. The low vacuity index and the no significant changes in %FI during the year emphasized the importance of the intertidal zone as a feeding ground for dog snapper.


Several studies have been carried out on the dietary habits of snappers (e.g. Hiatt and Strasburg, 1960; Guevara et al., 1994; Rooker, 1995; Kiso and Mahyam, 2003; Kamukuru and Mgaya, 2004; Szedlmayer and Lee, 2004). These studies suggested that Lutjanidae species were demersal predacious fishes with a broad diet dominated by crabs, shrimps, stomatopods, fish and other motile invertebrates.

According to the obtained data, the diet of L. jocu included a wide range of prey taxa related to the high diversity and abundance of food sources in the intertidal mangrove habitat. Similar results were reported for L.johnii(Bloch, 1792) in the Matang mangrove estuary (Malaysia) by Kiso and Mahyam (2003) and for L. campechanus (Poey, 1860) in Gulf of Mexico (USA) by Szedlmayer and Lee (2004).

According to Kamukuru and Mgaya (2004), while the range of prey consumed by L. jocu was large, comparatively few prey items dominated the diet. In this study, the analysis of stomach contents showed that the most frequent and dominant item was Penaeidae. In terms of biomass, the Grapsidae crabs presented a significant contribution in diet given the higher body mass. Our data are in agreement with the finding of Stark (1971) where juveniles of L. griseus (Linnaeus, 1758) from Florida, displayed a diet dominated by crustacea mainly crabs and shrimps (Penaeidae).

According to the %IRI values, Penaeidae, Grapsidae and Porcellanidae were the most important prey families of the juvenile dog snapper diet. Similar findings were reported for congeneric species such as the sub-adult of L. fulviflama(Forsskal, 1775) from Tanzânia showing a diet dominated by crustacea, with shrimps accounting for 40% of %IRI (Kamukuru and Mgaya, 2004).

During the sampling period, clear changes were observed in the diet composition of L. jocu. Dog snapper, reflected in the dried months a specialist diet, mainly consuming Penaeidae shirmps, and in the wetted months a generalist diet displaying a widening of the trophic spectrum consuming Grapsidae, Porcellanidae, Portunidae, Penaeidae and Xanthidae. Nevertheless, the occurrence of several prey items with lower contribution in abundance indicated that L. jocu was also an opportunistic predator. A possible cause for the observed intra-annual difference in diet of the juvenile dog snapper can be related to the distribution, abundance and availability of prey in each season (Guevara et al., 1994, Rooker, 1995, Sierra and Popova, 1997).



The authors thank M.F. Arnour, A. Jesus, E. Lameira and B. Almeida for assistance in field collections and laboratory analyses. This work was funded by the Millennium Initiative Project Coastal Resources ( financed by the Ministry of Science and Technology of the Brazil and the project MADAM (Mangrove Dynamics and Management) a cooperation between the Center for Tropical Marine Ecology (ZMT), Bremen, Germany, and the Universidade Federal do Pará (UFPA), Belém, Brazil. T. Giarrizzo acknowledges financial support by the Fundação de Amparo à Pesquisa do Estado do Pará (FAPESPA) (Project number: 137/2008 – Universal).



Amundsen, P. A.; Gabler, H. M. and Staldvik, F. J. (1996), A new approach to graphical analysis of feeding strategy from stomach contents data – modification of the Costello (1990) method. J. Fish Biol., 48, 607-614.         [ Links ]

ANA: Agência Nacional das Águas (2005), Accessed April 30, 2005. Available from:         [ Links ]

Cervigón, F. (1993), Los peces marinos de Venezuela. Volume II. Fundação científica de los Roques. Caracas, Venezuela.         [ Links ]

Cervigón, F.; Cipriani, R.; Fischer, W.; Garibaldi, L.; Hendrickx, M.; Lemus, A. J.; Márques, R.; Poutiers, J. M.; Robaina, G. and Rodriquez, B. (1992), Guia de campo de las espécies comerciales marinas y de aguas salobres de la costa septentrional de Sur America. Organizacion de las Naciones Unidas para la Agricultura y la Alimentacion. Rome.         [ Links ]

Cocheret de la Moriniére, E.; Pollux, B. Y. A.; Nagelkerken, I. and Van der Velde, G. (2002), Post-settlement life cycle migration patterns in relation to biotope preference of coral reef fish that use seagrass and mangrove habitats as nurseries. Estuar. Coast. Shelf Sci., 55, 309-321.         [ Links ]

Cocheret de la Moriniére, E.; Pollux, B. Y. A.; Nagelkerken, I. and Van der Velde, G. (2003), Diet shifts Caribbean grunts (Haemulidae) and snappers (Lutjanidae) and the relation with nursery-to-coral reef migrations. Estuar. Coast. Shelf Sci., 57: 1079-1089.         [ Links ]

Costello, M. J. (1990), Predator feeding strategy and prey importance: a new graphical analysis J. Fish Biol., 36, 261-263.         [ Links ]

Druzhinin, A. D. (1970), The range and biology of snappers (Fam. Lutjanidae). J. Ichthyol., 10, 715-735.         [ Links ]

Figueiredo, J. L. and Menezes N. A. (1978), Manual de peixes marinhos do sudeste do Brasil. II. Teleostei (1). Museu de Zoologia da Universidade de São Paulo-SP.         [ Links ]

Frédou, T. and Ferreira, B. P. (2005), Bathymetric trends of northeastern brazilian snappers (Pisces, Lutjanidae): Implications for the Reef Fishery Dynamic. Braz. Arch. Biol. Technol., 48, 787-800.         [ Links ]

Froese, R. and Pauly, D. (eds.) (2006), FishBase. Accessed January 30, 2006. Available from:         [ Links ]

Giarrizzo, T and Saint-Paul, U. (2008), Ontogenetic and seasonal shifts in the diet of the pemecou sea catfish Sciades herzbergii (Siluriformes: Ariidae), from a macrotidal mangrove creek in the Curuçá estuary, Northern Brazil. Rev. Biol. Trop., 56, 861-873.         [ Links ]

Giarrizzo, T. and Krumme, U. (2007), Spatial differences and seasonal cyclicity in the intertidal fish fauna from four mangrove creeks in a salinity zone of the Curuçá estuary, North Brazil. Bull. Mar. Sci., 80, 739-754.         [ Links ]

Giarrizzo, T. and Krumme, U. (2008), Heterogeneity in intertidal fish fauna assemblages along the world's longest mangrove area in northern Brazil. J. Fish Biol., 72, 773-779.         [ Links ]

Giarrizzo, T. and Krumme, U. (2009), Temporal patterns in recruitment of selected tropical fish to the mangrove creeks: implication for fisheries management in North Brazil. Braz. Arch. Biol. Technol., 52, 679-687.         [ Links ]

Guevara, C. E.; Bosch, M. A.; Suárez, M. R. and Lalana, R. R. (1994), Alimentación natural de tres especies de pargos (Pisces: Lutjanidae) en el Archipiélago de los Canarreos, Cuba. Rev. Investig. Marin., 15, 63-72.         [ Links ]

Hacunda, J. S. (1981), Trophic relationships among demersal fishes in a coastal area of the Gulf of Maine. Fish. Bull., 79, 775–788.         [ Links ]

Hiatt, R. W. and Strasburg, D. W. (1960), Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecol. Monogr., 30, 65-127.         [ Links ]

Hurlbert, S. H. (1978), The measurement of niche overlap and some relatives. Ecology, 59, 67-77.         [ Links ]

Isaac, V. N. (2006), Explotação e manejo dos recursos pesqueiros do litoral amazônico: um desafio para o futuro. Ciencia e Cultura, 58, 33-36.         [ Links ]

Kamukuru, A. T. and Mgaya, Y. D. (2004), The food and feeding habits of blackspot snapper, Lutjanus fulviflamma (Pisces: Lutjanidae) in Shallow waters of Mafia Island, Tanzania. Afr. J. Ecol., 42, 49-58.         [ Links ]

Kiso, K. and Mahyam, M. (2003), Distribution and feeding habitas of juvenile and yong John's snapper Lutjanus johnii in the Matang mangrove estuary, west coast of Peninsular Malaysia. Fish. Sci., 69, 563-568.         [ Links ]

Krebs, C. J. (1989), Ecological methodology. Harper Collins Publishers, New York.         [ Links ]

Krumme, U.; Keuthen, H.; Barletta, M. and Saint-Paul, U. (2005), Contribution to the feeding ecology of the predatory wingfin anchovy Pterengraulis therinoides (L.) in nothern Brazilian mangrove creeks. J. Appl. Ichthyol., 21, 469-477.         [ Links ]

Laegdsgaard, P. and Johnson, C. (2001), Why do juvenile fish utilise mangrove habitats? Journal of Exper. Mar. Biol. Ecol., 257, 229-253.         [ Links ]

Laegdsgaard, P. and Johnson, C. R. (1995), Mangrove habitats as nurseries: unique assemblages of juvenile fish in subtropical mangroves in eastern Australia. Mar. Ecol. Prog. Ser., 126, 67–81.         [ Links ]

Luckhurst, B. E.; Dean, J. M. and Reichert, M. (2000), Age, growth and reproduction of the lane snapper Lutjanus synagris (pisces: Lutjanidae) at Bermuda. Mar. Ecol. Prog. Ser., 203, 255-261.         [ Links ]

Marshall, S. and Elliot, M. (1997), A comparison of univariate and multivariate numerical and graphical techniques for determining inter- and intraspecific feeding relationships in estuarine fish.J. Fish Biol., 51, 526–545.         [ Links ]

Melo, G. A. S. (1996), Manual de identificação dos Brachyura (caranguejos e siris) do litoral brasileiro. Plêiade, São Paulo.         [ Links ]

Melo, G. A. S. (1999), Manual de identificação dos crustacea decapoda do litoral brasileiro: Anomura, Thalassinidea, Palinuridea e Astacidea. Plêiade, São Paulo.         [ Links ]

Menezes, N. A. and Figueiredo, J. L. (1980), Manual de peixes marinhos do sudeste do Brasil. IV. Teleostei (3). Museu de Zoologia, Universidade de São Paulo.         [ Links ]

Menezes, N. A. and Figueiredo, J. L. (1985), Manual de peixes marinhos do sudeste do Brasil. V. Teleostei (4). Museu de Zoologia, Universidade de São Paulo.         [ Links ]

Miranda, A. S.; Elorduy-Garay, J. F.; Villalejo-Fuerte, M. and Rojas-Herrera, A. A. (2003), Desarrollo gonadal y ciclo reproductivo de Lutjanus peru (Pisces: Lutjanidae) en Guerrero, México. Rev. Biol. Trop., 51, 489-502.         [ Links ]

Morton, R. M. (1990), Community structure, density and standing crop of fishes in a subtropical Australian mangrove area. Mar. Biol., 105, 385–394.         [ Links ]

Nagelkerken, I.; Van Der Velde, G.; Gorissen, M. W.; Meijer, G. I.; Van't Hof, T. and Den Hartog, C. (2000), Importance of mangroves, seagrass beds ands the shallow coral reef ar a nursery for important coral reef fishes, using a visual census technique. Estuar. Coast. Shelf Sci., 51, 31-44.         [ Links ]

Odum, W. E. and Heald, E. J. (1972), Trophic analyses of an estuarine mangrove community. Bull. Mar. Sci. 22, 671–738.         [ Links ]

Parrish, J. K. (1989), Re-examining the selfish herd: are central fish safer? Anim. Behav., 38, 1048–1053.         [ Links ]

Pinkas, L.; Oliphant, M. S. and Iverson, I. L. K. (1971), Food habits of albacore, bluefin tuna, and bonito in California waters. California Department of Fish and Game. Fish. Bull., 152, 1-105.         [ Links ]

Platell, M. E and Potter I. C. (2001), Partitioning of food resources amongst 18 abundant benthic carnivorous fish species in marine waters on the lower west coast of Australia. J. Exper. Mar. Biol. Ecol., 261, 31-54.         [ Links ]

Polunin, N. V. C. and Roberts, C. M. (1996), Reef fisheries. Fish Fish., 20, 477-485.         [ Links ]

Rezende S. M.; Ferreira B. P. and Fredou, T. (2003), A pesca de lutjanídeos no nordeste do Brasil: histórico das pescarias, características das espécies e relevância para o manejo. Bol. Téc. Cient. CEPENE, 11, 1-17.         [ Links ]

Rezende, S. M. and Ferreira, B. P. (2004), Age, growth and mortality of dog snapper Lutjanus jocu (Bloch and Schneider, 1801) in the northeast coast of Brazil. Braz. J. Ocean., 52, 107-121.         [ Links ]

Robertson, A. I. and Blaber, S. J. M. (1992), Plankton, epibenthos and fish communities. In- Tropical Mangrove Ecosystems (Coastal and Estuarine Studies 41), eds. Robertson, A. I. and Alongi, D. M. American Geophysical Union, Washinhton, DC, pp. 63-100.         [ Links ]

Rojas, L. P.; Maravilla, E. and Chicas, E. (2004), Habitos alimentarios del pargo mancha Lutjanus guttatus (Pisces: Lutjanidae) em los Cobanos y Puerto la libertad, El Salvador. Rev. Biol. Trop., 52, 163-170.         [ Links ]

Rooker, J. R. (1995), Feeding ecology of the schoomaster snapper Lutjanus apodus (Walbaum), from South-western Puerto Rico. Bull. Mar. Sci., 56, 881-894        [ Links ]

Serrano-Pinto, V. and Caraveo-Patiño, J. (1999), Survival of amarillo snaper Lutjanus argentiventris (Peters 1869) at different salinities in captivity. Aquac. Res., 30, 467-470.         [ Links ]

Sheaves, M. and Molony, B. (2001), Coherent patterns of abundance and size of tropical snapper in dynamic estuary systems. Wetl. Ecol. Manag., 9, 429-439.         [ Links ]

Sheaves, M. J. (1995), Large lutjanidae and serranidae fish in tropical estuaries: are they adults or juveniles?Mar. Ecol. Prog. Ser., 129, 31-40.         [ Links ]

Sierra, L. M. and Popova O. A. (1997), Relaciones tróficas de los juveniles de cinco especies de pargo (Pisces: Lutjanidae) en Cuba. Ver. Biol. Trop., 44/45, 499-506.         [ Links ]

Souza, R. F. C. (2002), Dinâmica populacional do pargo Lutjanus purpureus, Poey, 1875 (Pisces Lutjanidae) no Norte do Brasil. Master thesis, Universidade Federal do Pará, Pará, Brazil.         [ Links ]

Stark, W. A. (1971), Biology of the gray snapper, Lutjanus griseus (Linnaeus). Stud. Trop. Oceanogr., 10, 11–150.         [ Links ]

Szedlmayer, S. T. and Lee, J. D. (2004), Diet shifts of juvenile red snapper (Lutjanus campechanus) with changes in habitat and fish size. Fish. Bull., 102, 366-375.         [ Links ]

Underwood, a. j. (1997), Experiments in ecology. Their logical design and interpretation using analysis of variance. Cambrige University Press, Cambrige.         [ Links ]

Vazzoler, A. E. A. M. (1996), Biologia da reprodução de peixes teleósteos: teoria e prática. Eduem, Maringá         [ Links ].



Received: November 06, 2006;
Revised: November 30, 2007;
Accepted: June 12, 2009.



* Author for correspondence:

Creative Commons License All the contents of this journal, except where otherwise noted, is licensed under a Creative Commons Attribution License