Livestock Research for Rural Development 25 (9) 2013 Guide for preparation of papers LRRD Newsletter

Citation of this paper

Comparative analysis of diet of two sympatric species of Tilapia in Ayamé man-made lake (Côte d’Ivoire)

H Shep*, K M Konan, M Ouattara, A Ouattara and G Gourène

Laboratoire d’Environnement et de Biologie Aquatique, Université NANGUI ABROGOUA, 02 BP 801 Abidjan 02, Côte d'Ivoire   /
* Direction de l’aquaculture et des pêches (Ministère des Ressources Animales et Halieutiques),
Treichville, Rue des Pêcheurs BPV 19 Abidjan, Côte d'Ivoire


Tilapia zillii and T. guinensis, occurring in brackish and fresh waters of West Africa, respectively, are sympatric in artificial environment of the Ayamé Lake. The present study analyzed the stomach contents of 243 specimens of both species collected in Ayamé Lake(122 forTilapia zillii and 121 for Tilapia guineensis)in order to determine their food composition and interspecific diet overlap. Food items identified from stomach contents and food overlap were analysed using the Index of Food Preponderance (IFP) and the Morisita index modified by Horn.


The two species feed on a wide variety of items preys from animal and plant origin. T. zillii consumed 13 prey items while T. guinennsis feed on 16, among which 13items were common in their diets. The insect diptera and the macrophytes were the most important item in the diet of both species. Molluscs, coleoptera and zooplankton alsoconstituted some significant items. The food overlap analysis showed that T. zillii and T. guineensis share a wide range of prey types witha generalized feeding behaviour. Despite the wide food overlap, the competition for food is probably minimal, because the two fish species eat a wide range of foods and also because of the abundance of main preys.

Key words: artificial lake, diet composition, diet overlap, Tilapia guineensis, Tilapia zillii


Cichlid species are widely distributed in most east and west Afro-tropical hydrosystems (Daget 1988) and belong to the commercially important inland fishes of Africa (Fagade 1971). T. zillii is a well known species from West Africa river system through Chad basin to the Nile (Trewavas 1982, Negassa and Getahun 2004). The natural distribution of the species included Lake Albert, Lake Turkana, Israel and Jordan Valley (Trewavas 1982). T. zillii has been introduced throughout the world for production and vegetation control purposes (Spataru 1978). T. guineensis is a euryhaline species found in estuaries and lagoons of West Africa (Philippart and Ruwet 1982, Campbell 1987). This species has gained an increasing interest for aquaculture purposes, particularly in the estuaries and extensive lagoon systems which constitute its natural range (Akinwumi 2003, Akinrotimi 2006). T. zillii and T. guineensis occur in brackish and fresh waters of West Africa. T. zillii frequents the upper waters whereas T. guineensis is generally found in the lower parts and in the lagoons (Philippart and Ruwet 1982, Agnèse et al 1998). The artificial environment of the Ayamé Lake has favoured the coexistence of T. zillii and T. guineensis (Thys van den Audenaerde 1970). In other respects, a large population of hybrids of Tilapia species as results of hybridization among T. zillii and T. guineensis was found in Ayamé Lake (Pouyaud 1994).According to Pouyaud and Agnèse (1995), both species were genetically very close.Fisheries data from the lake revealed the predominance (52%) of tilapia species in the different landings sites around the Lake (Vanga et al 2002). Living in the same environment, these species likely share resources as well as reproduction habitats in Ayamé Lake. Sabagh and Carvalho-e-Silva (2008) indicated that the resource partition in a community is essential for the understanding of species interactions. In other respects, diets are a fundamental aspect of each niche, and it seems reasonable to assume that the structure of a community is based mostly on the way that food is shared among coexisting species (Sihand Christensen 2001).


 In Africa, the diets of tilapias species have been subject of several studies (e.g. Fryer et al 1955, Fagade 1971, 1978, Lauzanne 1988,Abdel-malek 1972,Akinwumi 2003, Winemiller and Kelso-Winemiller 2003,Negassa and Getahum 2004, Oso et al 2006,Shalloof and Khalifa 2009, Agbabiaka 2012). Particularly, T. zillii was considered as an opportunistic bottom feeder with a high proportion of the food consisting of plants parts (Nwadiaro 1984,Akinwumi 2003, Negassa and Getahun 2004), insects larvae, zooplankton, small crustacean and molluscs (Lauzanne 1988, Agbabiaka 2012) while T. guineensis mainly feed on algal filaments, diatoms, sand grain and unidentified organic material (Fagade 1971).


Nevertheless, feeding is one of the most important biological factors and its abundance and variety influence the structure and composition of fish populations (Aranha et al 2000). In addition, diet analysis of fishes allows us to understand their feeding strategy, their intra-or interspecific potential interaction (competition and predation) and indirectly indicate community energy flow (Ramirez-Luna et al 2008).


The present study aimed to understand the coexistence of the two species in a man-made lake. Food traits are discussed in the relation to the interspecific competition and dietary overlap of these fishes.

Material and methods

Study area


The Ayamé Lake (0° 20’ S and 36° 5’ E) is an artificial freshwater lake situated in the south-east region of Côte d’Ivoire. The lake was built in 1959 in the river Bia and has an area comprised between 87 and 194 km2 with a mean depth of 30 m. The lake level is subject to fluctuations depending on local rainfall and evaporation (Reizer 1967).The Ayamé Lake is deep and open water characterised by muddy substrate and a low transparency with an annual mean Secchi disk of 110 cm (Kouamélan et al 1999). Fishes were sampled at two main sites in the lake (Bakro and Ayamé).

Figure 1. Map of the Bia river showing the different sampling sites (•).

 Sampling and stomach contents analysis


A total of 243 specimens of T. zillii, T. guineensis were collected over twenty four months period in two sites (Figure 1) from August 1995 to September 1997 using two batteries of gill-nets with mesh size 10, 12, 15, 20, 25, 30, 35, 40 and 50 mm. Distribution per species is as follow: 122 for T. zillii and 121 for T. guineensis. Standard length (SL) of fish caught was measured to the nearest 0.1 cm using a measuring board. The standard length ranged from 60 to 200 mm for T. zillii, 100 to 170 mm for T. guineensis. The stomach was then removed and preserved in pillbox containing 5% formol for further examination in laboratory.


In the laboratory, the stomach contents of each specimen were placed in a Petri disk and aggregates were dispersed with a few drops of water and filtered through 1000, 500 and 100 µm mesh size before microscopic examination. The different prey taxa were sorted, counted and weighed to the nearest 0.0001g. For each stomach, the food items were identified to the lowest possible taxonomic level using descriptions and keys from various sources (Lindley 1970, 1975, Dejoux 1974, Elouard and Levêque 1977, Caratini 1985).


Data analysis


Calculation of indexes


The food of the two cichlids was assessed essentially by the relative frequency (RF) and relative dominance (RD) (King 1989, 1991, 1994) according to the formulae:



Where fi = frequency of item i; Fi = frequency of the nth item (sum of all fi); di = frequency of item i as dominant dietary; Di = frequency of nth dominant item (sum of all di). The RF and RD of all dietaries are then summed up to 100%. The RF is a modification of the occurrence method (Hyslop 1980) commonly used in fish dietary studies. The RF method over-emphasizes the importance of the numerous small items while RD method over-emphasizes the importance of large items. An index of food preponderance (IFP) was calculated as the mean of % RF and % RD. Items with IFP ≥ 10% were considered as primary dietaries while those with IFP values comprised between 1 to 10 % were considered as secondary. Items with IFP less than 1% were classified as incidental.


For food habits variation in relation to size, three classes (juveniles, sub-adults and adults) were determined as follows:

   Calculation of food overlapp


Food overlap between species has been calculated, using the overlap measure of Morisita (1959) as modified by Horn (1966).



Where S is the total number of food categories and Xi and Yi are the proportion of total diet of species X and Y taken from a given category of food i.


Feeding strategy


To assess the feeding strategy along the studied period, 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: 


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


Overall food composition


The results of the food items recorded are represented in table1. Tilapia zillii and T. guineensis feed on a wide trophic spectrum of food constituted by 16 and 13 items preys, respectively.The main food item of T. zillii were macrophytes (IFP = 15.90 %) and insects diptera (IFP = 11.62 %). The secondary preys (IFP = 1 – 10%) of this species were composed by molluscs, coleoptera, zooplankton, hymenoptera, ephemeroptera, mud, trichoptera and animal debris. The most common food item in the stomach of T. guineensis is the insect diptera (IFP = 13 %). Secondarily, T. guineensis fed on macropthytes, molluscs, coleoptera, mud, zooplankton, hymenoptera, trichoptera, ephemeroptera, animal debris and sand.

Table 1.  Overall food composition of Tilapia zillii and Tilapia guineensis in Ayamé Lake.


Items number

IFP, %

Tilapia guineensis (n = 121)

Tilapia zillii

(n = 122)








































































Animal debris










Undetermined preys





n = number of examined specimens, IFP = Index of food preponderance


Food similarity between the two species (Cλ = 0.98) was superior than 0.60.  A wide diet overlap between the both Tilapia species in Ayamé Lake was observed. T. zillii and T. guineensis feed on the same resources.


Food in relation to fish size

The figure 2 presented the preponderance index of the major food items, for the various size classes. All stages (juveniles, subadults and adults) of both species mainly fed on insects (IFP = 22 - 43.75 %), macrophytes (IFP = 16 – 33 %) and mollusks (IFP = 12 – 33 %). Zooplankton in juveniles of T. zillii (IFP = 11 %) and other preys (sand, animal debris, mud and undetermined preys) in juvenile (IFP = 11.52 %) and adults (IFP = 16.59 %) of T. guineensis also constituted some primaries preys. Only the feeding pattern of T. guineensis was significantly different between size classes (total χ2: 23.12, ddl = 4, P = 0.000104).

Figure 2. Food variation in relation to fish size of Tilapia zillii and Tilapia guineensis in the Ayamé Lake.

Seasonal variation in diet composition


The food composition in relation to season is presented in table2. In both seasons, insects (IFP = 37.69 – 44.62%) and mollusks (IFP = 14 – 18 %) constituted primaries preys for the two fish species. These preys were taken innearlythe same quantitiesby the studied fish. The specimens of Tilapia guineensis also mainly feed on fish (12 %) and other preys (IFP = 15 %) in wet season while in dry season these are the macrophytes and other preys which are also mainly consumed. In both seasons, the other primaries preys in the stomach of T. zillii are macrophytes. Zooplankton and arachnids were secondary and incidental preys, respectively, in the studied fish in all seasons.


The feeding pattern was significantly different between the studied species in wet season (total χ2 : 43.65, ddl = 13, P <  0.0001) and dry season (total χ2 : 58.242, ddl = 13, P = 0.000092).

Table 2.  Seasonal food composition of Tilapia zillii and Tilapia guineensis in Ayamé Lake


IFP, %

Tilapia guineensis

Tilapia zillii

Wet season

Dry season

Wet season

Dry season




































IFP = Index of food preponderance

Feeding strategy

Analysis of feeding strategy, based on the Amundsen’s method (figure 3), showed that for the two species, almost all the preys were located below the prey importance axis indicating that these fishes exploited a broad niche with a generalized feeding behaviour. Considering the prey importance, Tilapia zillii’s diet is mostly dominated by the items 14 (Macrophytes), 1 (Diptera), 11 (Molluscs) and 5 (Coleoptera). These preys were broadly consumed by all specimens of this species. The remaining items were eaten occasionally and in relative small amounts. For T. guineensis, diet was mostly based on 1, 14, 11 and 12 (Zooplankton) which tend to be dominant preys. The other items are occasionally and rarely consumed.

Figure 3. Feeding strategy of individuals of Tilapia zillii and Tilapia guineensis
in Ayamé Lake (see table 1 for item number).


Tilapia species have been reported to be plankton and deposit feeders (Adiase 1969,Fagade 1971, 1978, 1982, Fryer and Iles 1972,Fagade and Olaniyan 1973, Harbott 1975,Pauly 1976, Buddington 1979, Akinwumi 2003, Winemiller and Kelso-Winemiller 2003,Negassa and Getahum 2004, Oso et al 2006,Agbabiaka 2012).The food of the species covers a wide spectrum. The major prey items of Tilapia zillii and Tilapia guineensis in the Ayamé Lake were macrophytes, insects and molluscs parts. On that account, the feeding habits of these species were similar to those reported by Fagade and Olaniyan (1972) in the Lagos lagoon on Tilapia guineensis and Sarotherodon melanotheron and Fagade (1979) on Tilapia guineensis from Lekki lagoon.In addition, a wide variety of invertebrates are taken as food by Tilapia zillii andT. guineensis. According to Amundsen graphical, the studied fishes exhibited a general feeding strategy. Stomach content analysis based on the index of food preponderance showed that in Ayamé Lake, Tilapia zillii, T. guineensisfeed on a wide range of food organisms that makes them euryphagous feeding with a food base comprising both plants and animals.T. zillii has been variously classifiedas plankton feeders, higher plant and algae feeders or macrophagous as well as mud suckers (Fagade 1971,Brown and Colgan 1984, Negassa and Getahun 2004).Nevertheless, food of plant origin was the major component of the diet of T. zillii in this study. The same diet has been reported for T. zillii from Lake Victoria by Welcomme (1979), for fish from Lake Quarun by Abdel-Malek (1972) and from Nile canal by Khallaf and Alne-na-ei (1987).For that reason, T. zillii were brought to Florida (USA) in 1961 by the Florida Game and Freshwater Fish Commission to investigate it`s potential use as a biological weed control (Courtenay and Robins 1973).Because of its diet, covering a wide spectrum of food ranging from various types of plankton to invertebrate, T. zillii can be classified as an omnivorous. This characterization was reported for the same species by Spataru (1978) in Lake Kinneret (Israel).


Examination of the diet of Tilapia guineensis indicatedthat there was high index of food preponderance of Insects in their stomachs. Apart from the major food items, studied fishes also picked a variety of other food items. The wide variety of items occurring in the stomachs of T. zillii (13 items preys), T. guineensis (16 items preys) is an indication that they are non selective in feeding. In fact, 280 phytoplankton taxon (Ouattara et al 2000), 29 macroinvertebrates (24 Insect taxons) with a high abundance of MolluscPotadoma liberiensis (Diomandé and Gourène 2005) and 56 zooplankton taxons (Ouattara et al 2007) were identified in Ayamé Lake.In addition, about 10% of Lake Surface was occupied by the macrophytes Pistia stratiotes (Etien and Arfi 1996). Other macrophytes as Bacopa crenata, Ceratopteris cornuta, Scieria verrucosa, Commelina diffusa, Vigna campestris, Gongronerna latifolium) were present in the lake (Sankaré et al 1986).


The two species are able to use many sources of protein as food. Liem (1980) stated that teleost including cichlids are able to exploit more than one source. This ability to exploit different varieties of food makes T. zillii and T. guineensis as omnivorous fishes. The overlap measure Cλ (0.98) is near to 1. This result indicated a great food overlaps between species (Zaret and Rand 1971). So, competition for food becomes possible. However, direct competition seemed to be avoided to some extent as a result of great food availability in Ayamé Lake. In fact, Ouattara et al (2007) reported that the density of zooplankton in the Ayamé Lake was 116675ind/m3. In addition, the lentic conditions created by the lake are favorable to algal development. According to Spataru (1978), it is an important strategy for survival and an advantage over the species competing for a specific food item. The same observations were reported by Ahmed (2011) for the three sympatric species of Cyprinid fish larvae in Al-Huwaiza marsh (Southern Iraq). The similarity in ecological niche was also accompanied by some discrete differences in the selection of complementary food items. The difference noted in the diversity of supplementary food items of the two species may be an active and immediate response to interspecific competition or other habitat factors. T. zillii and T. guineensis were observed to be omnivorous, with highest frequency of occurrence of insect and macrophytes. Although some cichlids are known to feed entirely on fish scales (Fryer et al 1955). The low occurrence of fish scales in the stomachs of these specimens of tilapia species suggests that this type of prey is not important in the diet of tilapia species in the Ayamé Lake.


In conclusion there is a strong trophic-niche overlap between Tilapia guineensis and Tilapia zillii. However,the occurrence of overlap, even to a high degree,does not necessarily mean that competition is present, if the resource is not limited.


This work is a part of Ivorian-Belgian project VLIR / KUL (Flemish Interuniversity Raad) " Évolution de la biodiversité des poissons après la construction d’un barrage: cas de la rivière Bia en Côte d’Ivoire" financed by the General Agency for Development Cooperation Bestuur voor Algemeen Ontwikkelingssamenwerking (ABOS-BADC) of Belgium. We would be grateful to the promoter D.F.E. Thys van den Audenaerde and the co-promoter, late G.G. Teugels.  


Abdel-malek SA 1972 Food and feeding habits of some Egyptian fishes in Lake Quarun: Tilapia zillii (Gerv.). B. According to different length groups. Bulletinof the Institute of Oceanography and Fisheries, Cairo. Volume 2:203-213.


Adiase K 1969 A preliminary report on the food of fish in the Volta Lake. In: Obeng LE (ed.), Man-made lakes. Ghana University Press, Accra. pp235-237.


Agbabiaka L A 2012 Food and feeding habits of Tilapia zillii (Pisces: Cichlidae) in river Otamiri South-eastern, Nigeria. BioscienceDiscovery. Volume 3(2): 46-148.


Agnèse J F, Adépo-Gourène B and Pouyaud L 1998 Natural hybridization in tilapias. In: Agnèse JF (Ed.) Genetic and Aquaculture in Africa, Paris, Edition ORSTOM,pp95-104.


Ahmed S M 2011 Note on feeding relationships of three species of cyprinid fish larvae in Al-Huwaiza marsh, Southern Iraq. Mesopotamian Journalof Marine Science. Volume 26 (1): 35-46.


Akinrotimi O A 2006 Effects of acclimation on haematoligical characteristics of black-chin tilapia.  Post Graduate Diploma Project, Department of Fisheries, Rivers State University of Science and Technology, Port Harcourt.pp65.


Akinwumi F O 2003 Food and feeding habits of Tilapia zillii (Pisces: Cichlidae) in Ondo State University Fish Farm. The 16th Annual Conference of FISON, 4-9 November 2001, Maiduguri, Nigeria. pp195-198.


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 Costello method. Journal of Fish Biology.Volume 48: 607-614.


Aranha J M R, Gomes J H C and Fogaça F N O 2000 Feeding of two sympatric species of Characidium, C. laneiand, C. pterostictum (Characidiinae) in a coastal stream of Atlantic Forest (Southern Brazil). Brazilian Archives of Biologyand Technology. Volume 43 (5): 527-531.


Brown J A and Colgan P W 1984 The ontogeny of feeding behaviour in four species of centrarchid fish. Behavioural Processes.Volume 9: 395-411.

Buddington R K 1979 Digestion of an aquatic macrophyte by Tilapia zillii (Gervais). Journal of Fish Biology.Volume 15: 449-455.


Campbell D 1987 A review of the biology and culture of Tilapia guineensis. African Regional Aquaculture Center, Port Harcourt, Nigeria.pp20.


Caratini R 1985 Zoologie. Editions Bordas, Paris.pp25-45.


Costello M J 1990 Predator feeding strategy and prey importance: a new graphical analysis. Journal of Fish Biology.Volume 36(2): 261-263.


Courtenay W R Jr and Robins C R 1973 Exotic aquatic organisms in Florida with emphasis on fishes: a review and recommendations. Transactions of the American Fisheries Society.Volume102 (1): 1-12.


Daget J 1988 Systématique. In: Lévêque C, Bruton MN and Ssentongo GW (eds.), Biologie et Ecologie des poissons d’eau douce africains ORSTOM. Coll. Trav. Doc. Paris, 216. pp15-34.


Dejoux C 1974 Contribution à la connaissance des Chironomidés de l’Afrique de l’Ouest. Entomologisk Tidskrift. Volume95: 72-83.


Diomandé D and Gourène G 2005 Premières données sur la macrofaune benthique de l’hydrosystème fluvio-lacustre de la Bia (Côte d’Ivoire). Sciences et Nature. Volume 2 (2) : 167- 176.


Duponchelle F and Legendre M 2000 Oreochromis niloticus (Cichlidae) in Lake Ayamé, Côte d’Ivoire: Life history traits of a strongly diminished population. Cybium. Volume24 (2): 161-172.


Duponchelle F and Panfili J 1998 Variation in age and size at maturity of female tilapia, Oreochromis niloticus, population from man-made lakes of Côte d’Ivoire. Environmental Biology of Fishes. Volume 52 (4): 453-465.


Elouard J M and Levêque C 1977 Rythme nycthéméral de dérive des insectes et des poissons dans les rivières de Côte d’Ivoire. Cahier ORSTOM, Série Hydrobiologie. Volume 11 (2): 179-183


Etien N and Arfi R 1996 Macrophytes aquatiques dans les eaux« continentales»Ivoiriennes. Archives Scientifiques, Centre de Recherche Océanologique, Abidjan. Volume XV (2) : 1-30.  or


Fagade S O 1971 The food and feeding habits of Tilapia in Lagos Lagoon. Journal of Fish Biology. Volume3: 151-156.


Fagade S O 1978 On the biology of Tilapiaguineensis from Lekki lagoon, Lagos State, Nigeria. Nigerian journal of science. Volume12: 73-87.


Fagade S O 1979 Observations on the biology of two species of Tilapia from the Lagos lagoon, Nigeria. Bulletin de l’Institut Fondamental d’Afrique Noire.Volume41: 627-653.


Fagade S O 1982 The food and feeding habits of Sarotherodon galilaeus from a small lake. Archiv fur Hydrobiologie. Volume93 (2): 256-263.

Fagade S O and Olaniyan C I O 1973 The food and feeding interrelationship of the fishes of Lagos lagoon.Journal of Fish Biology. Volume5: 205-227.


Fryer G, Greenwood P H and Trewavas E 1955 Scale eating habits of African cichlid fishes. Nature.Volume175: 1089-1090.


Fryer G and Iles T D 1972 The cichlid fishes of the great lakes of africa. Their biology and evolution. Oliver & Boyd, Edinburgh, pp641.


Harbott J 1975 The feeding biology of Sarotherodon niloticus in lake Turkana (Rudolf). In: Papers presented at the symposium on the hydrobiology and fisheries of Lake Rudolf. Molo


Horn H 1966 Measurement of overlap in comparative ecological studies. The American Naturalist. Volume 100: 419-424.


Hyslop E J 1980 Stomach contents analysis. A review of methods and their application. Journal of Fish Biology. Volume 17(4): 411-429.


Khallaf E A and Alne-na-ei A A 1987 Feeding ecology of Oreochromis niloticus (Linnaeus) &Tilapia zillii (Gervais) in a Nile Canal. Hydrobiologia.Volume 146: 57-62


King R P 1989 Distribution, abundance, size and feeding habits of Brienomyrus brachyistius (Gill, 1862) (Teleostei: Mormyridae) in Nigeria rainforest stream. Cybium. volume 13(1): 25-36.


King R P 1991 Some aspects of the reproductive strategy of Ilisha africana (Bloch, 1795) (Teleostei, Clupeidae) in Qua Iboe estuary,Nigeria. Cybium.Volume 15(3): 239-257.


King R P 1994 Seasonal dynamics in the trophic status of Papyrocranus afer (Günther, 1868) (Notopteridae) in a Nigerian rainforest stream.Revued’Hydrobiologie Tropicale. Volume 27(2): 143-155.


Kouamélan E P, Teugels G G, Gourène G, Ollevier F and Thys van den Audenaerde D F E 1999. The effect of man-made lake on the diet of African electric fish Mormyrus rume Valenciennes, 1846 (Osteoglossiformes; Mormyridae). Hydrobiologia. Volume 380 : 141-151.


Lauzanne L 1988 Les habitudes alimentaires des poissons d’eau douce africains.In: Lévêque C, Bruton MN et Ssentongo GW (eds.),  Biologie et Ecologie des poissons d’eau douce africains. ORSTOM Paris.pp395-425.


Liem K F 1980 Adaptive significance of intra and interspecific differences in the feeding repertories of cichlid fishes. American Zoologist.Volume 20: 295-314.


Lindley R P 1970 On a new genus and species of Libellulid dragonfly from the Ivory Coast. Entomologist London.Volume 103: 77-83.


Lindley R P 1975 The dragonfly of Korhogo, Ivory Coast. Bulletin de l’Institut Fondamental d’Afrique Noire.Volume 36(3): 682-698.


Morisita M 1959 Measuring of interspecific association and similarity between communities. Memoirs of the Faculty of Science Kyushu University Series E, 3: 65-80.


Negassa A and Getahun A 2004  Food habits and diel feeding rhythm of introduced fish, T. zillii Gervais, 1948 (Pisces: Cichlidae) in Lake Zwai, Ethiopia. SINET:Ethiopian Journal of Science. Volume 27(1): 9-16.


Nwadiaro C S 1984 A comparative study of the food habit and distribution of Tilapia Sarotherodon and Pelvicachromis in River Sombreiro, Nigeria.Revue de Zoologie et de Botanique Africaines. Volume 91 (1): 41-46.


Oso J A, Ayodele I A and Fagbuaro O 2006 Food and Feeding Habits of Oreochromis niloticus (L.) and Sarotherodon galilaeus (L.) in a Tropical Reservoir. World Journal of Zoology.Volume 1(2): 118-121.


Ouattara A, Podoor N, Teugels G G and Gourène G 2000 Les microalguesde deux cours d’eau (Bia et Agnébi) Côte d’Ivoire. Syst. Geogr. Pl.Volume 70 : 315-372.


Ouattara I N, Ouattara A, Koné T, N'Douba V and Gourène G 2007 Distribution du zooplancton le long de deux petits bassins côtiers ouest africains (Bia et Agnebi ; Côte d'Ivoire). Agronomie Africaine.Volume 19 (2): 197-210.


Pauly D 1976 The biology, fishery and potential for aquaculture of Tilapia melanotheron in a small West African lagoon. Aquaculture.Volume 7: 33-49.


Philippart J C l and Ruwet J C l 1982 Ecology and distribution of tilapias. In: Pullin R S V and Lowe-McConnell R H (eds.). The Biology and Culture of Tilapias. Proceedings of the international conference, ICLARM, Manilla, Phillipines.pp15-59.


Pouyaud L 1994 Génétique des populations de tilapias d’intérêt aquacole en Afrique de l’Ouest. Relations phylogénétiques et structurations populationnelles. PhD Thesis, University of Montpellier II, Sciences et technique du Languedoc. pp229.


Pouyaud L and Agnèse J F 1995 Phylogenetic relationships between 21 species of three tilapiine genera Tilapia, Sarotherodon and Oreochromis using allozyme data. Journal of Fish Biology. Volume 47: 26-38.


Ramirez-LunaV, Navia A F and Rubio E A 2008 Food habits and feeding ecology of an estuarine fish assemblage of northern Pacific coast of Equator. Pan-American Journal of Aquatic Sciences, Volume 3 (3): 361-372.


Reizer C 1967 Aménagement piscicole du lac artificiel d’Ayamé. Publication n°30 du Centre technique forestier tropical. pp 108.


Sabagh L T and Carvalho-e-Silva A M P T 2008 Feeding overlap in two sympatric species of Rhinella Anura (Bufonidae) of the Atlantic Rain Forest. Revista Brasileira de Zoologia.Volume 25 (2): 247-253.


Shalloof K A Sh and Khalifa N 2009 Stomach Contents and Feeding Habits of Oreochromis niloticus (L.) From Abu-Zabal Lakes, Egypt. World Applied Sciences Journal. Volume 6 (1): 01-05.


Sih A and Christensen B 2001 Optimal diet theory: when does it work, and when and why does it fail? Animal Behaviour. Volume 61: 379-390.

Spataru P 1978 Food and feeding habits of Tilapia zillii (Gervais) (Cichlidae) in Lake Kinneret (Israël). Aquaculture.Volume 14: 327-338.


Thys van den Audenaerde D F E 1970 An annoted bibliography of Tilapia (Pisces: Cichlidae). Documentation Zoologique (Musée Royal de l’Afrique Centrale). Volume 14. p 406.


Trewavas E 1982 Taxonomy and speciation. In: Pullin RSV and Lowe-McConnell RH (eds.). The Biology and Culture of Tilapias. Proceedings of the international conference, ICLARM, Manila, Philippines. pp3-13.


Vanga A F, Gourène G and Ouattara M 2002 Impact de la pêche sur la disposition en poisson dans les régions des lacs d’Ayamé et de Buyo (Côte d’Ivoire). Archives Scientifiques,Centre de Recherche Océanographique, Abidjan.Volume 22 (2): 1-12.


Welcomme R L 1979 Fisheries Ecology of Floodplain Rivers. Longman, London, ISBN-13: 978058246310. pp317.


Winemiller K O and Kelso-Winemiller L C 2003 Food habits of tilapiine cichlids of the Upper Zambezi River and floodplain during the descending phase of the hydrologic cycle. Journal of fish biology.Volume 63: 120-128.


Zaret T M and Rand A S 1971 Competition in tropical stream fishes: Support for the competitive exclusion principle. Ecology.Volume 52: 336-342.

Received 12 June 2013; Accepted 21 August 2013; Published 4 September 2013

Go to top