Fisheries Research 35 1998.

209218

Studies on aspects of the ecology of the catfish Chrysichthys auratus Geoffrey St. Hilaire Osteichthyes; Bagridae / in the River Benin Niger Delta, Nigeria /
R.B. Ikomi ) , O. Odum
Zoology Department, Delta State Uniersity, Abraka, USA Accepted 2 December 1997

Abstract Data on some aspects of the ecology of the catfish Chrysichthys auratus in the Benin river are presented. Fish were captured throughout the year. C. auratus constituted about 63% of the Bagrid catch and favoured the shallow littoral river zone with silty sandy substratum. It attained a total length of 14.6 cm with a weight of 32.0 g. The growth in weight relative to length was isometric and the condition factor ranged from 1.02.9. It increased with individual size of fish irrespective of sex and was higher during the wet season than in the dry season. C. auratus fed predominantly in insects, fish and aquatic macrophytes. Crustaceans and algae were a minor part of the diet. It was essentially a night feeder with no clear qualitative variation in feeding habit with size and season. Breeding occurred between May and July and estimates of egg number ranged from 260 to 620. Gonadosomatic index GSI. values ranged from 2.69 to 19.68. q 1998 Published by Elsevier Science B.V. All rights reserved.
Keywords: Chrysichthys auratus; Bagridae; Distribution; Food; Feeding habit; Growth pattern; Fecundity

1. Introduction The fish family Bargridae occurs principally in Tropical Africa Jayaram, 1966., and the genus Chrysichthys is widely distributed and of great economic importance in West Africa Welma, 1948; Jayaram Op. Cit.. In Nigeria, many investigations on the biology of catfishes of this genus have been conducted Imevbore and Okpo, 1975; Ezenwa, 1971; Ajayi, 1972; Ikusemiju and Olaniyan, 1975.. However, despite the abundant data on Chrysichthys,

Corresponding author.

there are very few biological studies on Chrysichthys auratus. This species has recently become important to the riparian communities of the Niger Delta area of Nigeria principally because of its affordability, tasteful flesh and relative abundance in comparison with other bagrids in the river. Published information is mainly taxonomic Lowe-McConnell, 1972; Anthony, 1982. or briefly descriptive of the biology Imevbore, 1970; Imevbore and Bakare, 1970; Lauzanne, 1972; Odum, 1992.. To date, information on the ecology of C. auratus in rivers of the Niger Delta is scarce. This paper therefore make some ecological observations on the species in the River Benin.

0165-7836r98r$19.00 q 1998 Published by Elsevier Science B.V. All rights reserved. PII S 0 1 6 5 - 7 8 3 6 9 8 . 0 0 0 7 1 - X

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2. Description of study site The River Benin 5835X 5835X E; 68:35X N; Fig. 1. is one of four major rivers that drain the Niger Delta basin. It runs through a swampy terrain at Koko and Sapele to its Atlantic estuary. The river bifurcates at Sapele where it is joined by two major tributaries, the Rivers Jamieson and Ethiope. The study site, lying within the evergreen forest zone, is characterized by freshwater and mangrove swamp vegetation. Common fringing freshwater trees are Mitragyna ciliata Aubreville pellegr; Grewia spp., Rhizophora harrisonii Leechman and Aeciennia africana P. Beauv. Two climatic seasons, wet and dry, prevail in this area. The wet season MayOctober. is characterized by the prevalence of the moist southwesterly wind from the Atlantic Ocean and high rainfall monthly average ) 20 cm.. The dry season NovemberApril. is characterized by low precipitation 020 cm. accompanied by the hot and dry northeastern wind from the Sahara desert. Variations in some key physicochemical parameters of the river are pH 67.4., conductivity 1127 US cmy1 ., salinity 13.5 mg ly1 . and alkalinity 23.5 mg

CaCO 3 ly1 .. Relevant human activities in the sites are logging, domestic washing and fishing.

3. Materials and methods Monthly sampling of fish in river Benin was conducted from three stations Fig. 1. between March 1994 and February 1995. Fish were captured during both night and day using three methods: set gill nets of 24.0 cm mesh size, hook and line baited with worms, and non-return valve wire straps. Day sampling was carried out between 0700 and 1200, while night sampling was between 2300 and 0500. Fish on capture were immediately preserved in 10% formalin solution prior to laboratory examination. In the laboratory, each fish was weighed to the lowest 0.1 g and the total length measured to the nearest 0.1 cm. Stomachs were removed and the contents identified to the lowest convenient taxon. Stomach contents were analyzed by the points and the frequency of occurrence methods Hynes, 1950; Hyslop, 1980.. In the point method, each non-empty stomach was assigned 20 points which were shared among the different contents of the stomach taking into account

Fig. 1. Map showing the location of River Benin in Nigeria inset. and map of River Benin showing sampling sites.

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their relative proportion by volume. The mean point scored for each food item was determined and expressed as a percentage of the total points accumulated across all items recovered from all stomachs examined. In the frequency of occurrence method, the occurrence of each food item was expressed as a percentage of the total number of stomachs containing food. The lengthweight relationship of the fish was described by the equation: w s al b where w is weight in grams, l is total length in centimeters and a and b are regression constants. The logarithmic transformed data gave the linear regression equation: Log w s log a q b log l The Fultons condition factor of each specimen was calculated from the relationship: 100W Ks 3 l where K is the condition factor. l is total length in centimeters and W is weight in grams. Sex was determined by visual and microscopic examination of the gonads. The unsexed small fishes were regarded as immature. Fecundity was estimated from ripe ovaries stage III. by direct count. The relationship between fecundity and fish lengthrweight was described by the equation: F s aX b where F is fecundity, X is body length in centimeters or body weight in grams, b is the slope and a the intercept Lagler, 1978.. Through a logarithmic transformation the equation becomes: Log F s log a q b log X The maturity stages of the ovary were classified using the modified Kestevens scheme Bagenal and Braun, 1978. as shown below: Stage I: immature ovary greyfish, thin and elongated. Eggs invisible to naked eyes.. Stage II: ripening ovary light yellow, eggs visible through ovary wall, oocyte diameter 0.61.5 mm.. Stage III: ripe ovary bright yellow, full and swollen. All eggs distinct, round and fully yolked. Egg diameter 1.62.1 mm.. Stage IV: ripe running ovary as in stage III but slightly flaccid with numerous deep yellow coloured eggs. Some translucent. Egg diameter 1.82.1 mm..

Stage V: spent ovary flaccid, opaque coloured and shrunken. Primary oocyte in stage I. Considerable atresia..

Fig. 2. Length-frequency distribution of Chrysinchthys auratus from the River Benin.

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The gonadosomatic index GSI. was calculated for each gonad from the equation GSI s Gonad weight g . Fish weight g . = 100

dentalis Valenciennes and Auchenoglanis biscutatus Geoffrey St. Hilaire. C. auratus constituted about 63% of the total bagrid collected and no other species exceeded 13%, thus C. auratus was the dominant bagrid in the river. 4.2. Lengthweight relationship The weight of C. auratus sampled ranged from 1.6 to 32.0 g. The least squares common fit analysis of the transformed data for each sex and for combined sexes pooled resulted in the equations in Table 2. The regression slopes indicated an approximately isometric growth pattern. A scatter plot of the total length and body weight relationship of the combined sexes is shown in Fig. 3. The figure reflects the exponential growth in weight with increasing length. 4.3. Condition factor The mean monthly condition factor declined continuously from 2.91.2 throughout the MayOctober wet season and until November. It then increased in the dry season to its May peak Fig. 4.. Overall K-values increased with individual size of fish irrespective of sex Fig. 5.. However, they were slightly higher in the males than in the females. 4.4. Food and feeding Stomach fullness, stomach content and dietary composition of 464 C. auratus whose length ranged from 2.4 to 14.6 cm were examined. Four hundred and forty-one had food in their stomach and 23 had empty stomachs Table 3.. The fish fed primarily on

The Students t-test parameter to compare calculated slope in lengthweight regressions and the chi-square test evaluating diet variations in food habit of C. auratus were adopted from Zar 1984..

4. Results 4.1. Distribution and abundance C. auratus captured in the River Benin ranged from 2.4 to 14.6 cm total length. The largest male measured 14.6 cm and female 13.8 cm. The monthly length frequency distribution shows several modes Fig. 2. which cannot be said to represent definite year classes. Recruitment of the juveniles occurred in most months of the year. Adult specimen F 5 cm. and immature specimens - 5 cm. were captured throughout the study period. C. auratus was ubiquitous in the river but both the adults and juvenile prefer shallow portions of the river - 1.5 m. with silty sandy substratum. The species was most active at night and occurred more frequently in night catches irrespective of the gear used Table 1.. The gill net appears to be the most effective sampling gear. It captured over 60% of the C. auratus used in this study. During sampling, 739 bagrids were captured: Chrysichthys nigrodigitatus Lecepede, Chrysichthys auratus Geoffrey St. Hilaire, Auchenoglanis occi-

Table 1 Variations in the number of C. auratus captured using the three different sampling gears at both day and night Gears time interval h. Day haul 7.0012.00 Night haul 23.005.00 Total number Percentage Gill net 111 188 299 64.4 Wire traps 42 76 118 25.4 Hook and line 16 31 47 10.2

R.B. Ikomi, O. Odum r Fisheries Research 35 (1998) 209218 Table 2 Lengthweight relationship in C. auratus Size total length. cm. 2.45.0 5.114.6 5.113.8 2.414.6 Individual immature male female combined sex Lengthweight relation log log log log w s y1.84 q 2.97 log w s y1.89 q 3.00 log w s y1.81 q 2.98 log w s y1.89 q 2.99 log l l l l N 96 176 192 464 r 0.688 0.731 0.791 0.712 t 0.76 1.41 1.38 1.48 P ) 0.05 ) 0.05 ) 0.05 ) 0.05

213

n s paired number, r s correlation coefficient, t s absolute value of the t-test parameter to compare calculated slope to 3.

insects, fish, and aquatic macrophytes with algae, crustaceans and molluscs of secondary importance. The insects which were the most important food item consisted mainly of dipteran and trichopteran larvae. Aquatic macrophytes, mainly plant seeds and leaves, were also heavily consumed. Other food items were taken only sparingly. Additionally, sandgrains occurred in appreciable quantities in the stomach of the fish. Fig. 6 shows the stomach content of C. auratus in relation to length class. The figure clearly demonstrates changes in diet with size particularly with the minor parts of the diets. For example, with increasing length, the fish consumed proportionally more fish, molluscs and macrophytes while the amount of algae and crustaceans decreased. Both the small and large C. auratus fed heavily on insect larvae.

Information on the temporal diet variation is also shown in Table 3. Quantitatively, in both the wet and dry seasons, C. auratus fed on all the food items listed in Table 3 except bivalve molluscs which were not taken during the wet season. However, quantitatively, insects and fish contributed more during the wet season. On the other hand, more crustaceans copepods, and shrimps. and macrophytes were consumed during the dry season. The other food items were taken in variable quantities. Also, the different groups of food items consumed during the wet season numbered 9 while the dry season figure was 10 Table 3.. The difference in the number was not significant x 2 s 0.052; P ) 0.05.. The diel food pattern was investigated using all the fish captured. One hundred and eighty-nine were captured during the day and 275 at night. Of the day sample, 173 had food in their stomach while among the night specimens, the corresponding figure was 268. The night time specimens had significantly

Fig. 3. Scatter plot of the lengthweight relationship of C. auratus.

Fig. 4. Variations in the mean monthly condition factor of C. auratus.

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Fig. 5. Variation in the relative condition factor in the different length classes on C. auratus.

more food x 2 s 30.0; P - 0.001. than the day ones. Copepods and molluscs were not fed on at night Table 4. and were of secondary importance. Other food items listed were consumed by both day and night. The occurrence of the primary diet items fish and insects. was significantly higher P - 0.001. at night. However, the occurrence of aquatic macrophytes in the diet was significantly greater in the day than at the night. Also, the fluctuations in the points scored for the various food items followed the same pattern as in the occurrence method except for the insects where no significant difference P ) 0.05. occurred between the day and night Table 4..

Fig. 6. Variations in the feeding habit in the different length classes of C. auratus.

4.5. Reproduction The numbers of the males and females recorded were 196 and 192, respectively. The observed male:female ratio of 1:0.98 was not significantly different from the expected ratio of 1:1 x 2 s 0.04;

Table 3 Gross tropic spectrum and seasonal variation in the percentage occurrence %O. and percentage point %P. of the diets of C. auratus in River Benin Food items Gross composition %O Fish Dipteran larvae Trichopteran larvae Insect remains Macrophyta Algae Copepoda Malacostraca shrimps. Mollusca Sand No. of fish examined No. of empty stomachs No. of categories of food items 17.2 44.3 39.4 20.1 16.3 8.0 3.7 3.2 5.1 40.1 464 23 10 %P 23.7 15.3 10.4 15.4 8.7 4.0 1.3 4.0 3.6 13.6 Wet season %O 17.6 27.2 10.1 12.3 5.2 8.6 6.2 12.2 0 21.7 284 7 9 %P 33.1 22.5 10.2 8.2 6.3 3.0 2.4 8.6 0 5.7 Dry season %O 8.5 29.3 12.1 3.6 14.3 9.2 8.9 13.7 6.3 21.6 180 16 10 %P 10.4 20.2 11.3 7.9 10.3 3.0 5.6 15.1 7.3 8.9

R.B. Ikomi, O. Odum r Fisheries Research 35 (1998) 209218 Table 4 Diel variation in food habits of C. auratus from River Benin Methods Occurrence Day Fish Dipteran larvae Trichopteran larvae Insect remains Macrophyta Algae Copepoda Malacostraca Shrimps. Mollusca Sand 7.9 34.9 34.4 10.6 33.3 13.8 9.0 1.6 12.7 30.7 Night 23.6 50.9 42.9 30.2 9.1 5.5 0 4.4 0 5.8 Point

215

x2
7.83 2.98 0.95 9.42 13.8 3.87 y 1.3 y 16.99

P - 0.001 0.001 ) 0.05 - 0.001 - 0.001 - 0.001 y 0.05 - 0.001

Day 16.5 10.6 8.8 11.7 17.5 7.8 3.9 2.9 10.6 9.7

Night 32.3 15.0 12.1 16.8 5.2 2.3 0 5.2 0 11.1

x2
5.12 0.76 0.52 0.91 6.66 2.99 y 0.65 y 0.89

P - 0.001 ) 0.05 ) 0.05 ) 0.05 - 0.001 ) 0.05 y ) 0.05 y ) 0.05

The values shown for day and night are percentage frequencies and points gained.

P - 0.001.. Using the Kesteven scheme, five gonadal maturity stages were recorded Table 5.. Mature females with ripe ovaries stage III. were caught in most months of the year, however, a major percentage occurred in May. Ripe running females stage IV. dominated the catch in June while spent females stage V. were encountered mostly in JulyrAugust. Immature fishes were captured throughout the period under investigation but occurred most in September to November. Ripening ovaries were recorded principally between January and April.

The fecundity estimate varied with size of fish and the average number of eggs in a ripe ovary ranged from 260 to 620 within the size range of 10.214.0 cm, total length Fig. 7.. The minimum and maximum numbers were counted in specimens with total length 10.5 and 13.8 cm respectively.

Table 5 Percentage frequency distribution of the female gonad maturity stages and mean gonadosomatic index GSI. of C. auratus during sampling period Months Stages I March April May June July August September October November December January February 1994 1994 1994 1994 1994 1994 1994 1994 1994 1994 1995 1995 5 10 13 5 17 9 58 56 44 11 23 14 II III IV V 47 48 8 15 8 18 25 26 21 45 53 57 21 24 59 20 8 9 8 6 14 22 8 19 16 18 16 50 26 18 8 6 14 11 8 10 11 8.9 0 11.3 4 14.6 10 19.6 42 3.7 46 3.2 0 2.69 6 2.69 7 3.0 11 3.61 8 6.31 0 6.91 19 21 24 20 12 11 12 16 14 9 13 21 Fig. 7. Variations in the fecundity values of Chrysichthys auratus in relation to length. GSI %. Total examined

Stages: I, immature; II, ripening; III, ripe; IV, ripe running; V, spent.

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The logarithmic transformed data of the lengthrweight and fecundity relationship provided the results below: Log F s 1.7406 q 0.7903 log l r correlation coefficient. s 0.294; N s 48 Log F s 2.3784 q 0.1595 log W r s 0.1723; n s 48 Fecundity was not significantly correlated P ) 0.05. either to total length or body weight. The ripe eggs were yellowish in colour. Egg diameters ranged from 1.62.1 mm with a mean of 2.0 mm. Eggs in all ovaries were generally at similar stages of development. The mean monthly GSI values ranged between 2.69 and 19.6% Table 5.. The GSI values were greater in May and June ) 14.0%. but dropped drastically to less than 4.0% between July and December. Thereafter, there was gradual rise in GSI values.

5. Discussion Chrysichthys auratus evidently is the ecologically dominant bagrid in River Benin. Its dominance among the bagrids of Lake Kainji in Nigeria has been reported by Ajayi 1972.. Adult and juvenile C. auratus flourished in all stretches of the river but showed preference for the shallow littoral portions. This behavioural adaptation may have been partly influenced by the distribution of the aquatic insects which constituted food items. According to LoweMcConnell 1975., such littoral zones provide thriving habitats for most aquatic insects. Additionally, this behaviour probably protects them against large predators usually encountered in the deeper portions of the river. C. auratus appears to be more active at night than by day as indicated by the higher number of specimens captured in the night sampling. In the river, it did not attain large size. Its maximum length was 14.6 cm as against a record length of 31.1 cm obtained for the species in the Kainji Lake, Nigeria, by Bakare 1968.. The length frequency plot showed several modes which may have resulted from different broods irregularly produced at various months of the year. The regression slopes b-values obtained for weight and length relationship were not significantly different from 3, indicating

that the species exhibits isometric growth Bagenal and Tesch, 1978.. The condition factor varied with the month and appears to be influenced by the rainfall regime and breeding activities of the fish. During the wet season, mean K-values were relatively higher than in the dry season. This probably resulted from the effective utilization of the relatively rich rainy season resources of the river personal observation.. In addition, the month of May recorded higher K-value and this coincided with the period in which the most mature adult carried ripe gonads. The sudden drop in K-value in June probably reflects fish spawning activities Table 5.. In both males and females mean K-values improved with increase in individual length of fish. It is possible that its foraging ability and conservation of stored food energy improves with increasing size; it is also possible that increasing weight of maturing gonads may be partially responsible for the observed increase in K-values with fish length. C. auratus is versatile in food habit and this is reflected in the high percentage ) 95%. of stomachs containing food. Food items of animal origin contributed significantly to the diet, which was dominated principally by fish Tilapia and Brycinus spp., bottom dwelling immature dipteran mainly Chironomus and Chaoborus., and trichopteran larvae. In addition, vegetable matter contributed significantly. This helps to meet the carbohydrate need of the fish. C. auratus in the River Benin is an omnivore. This substantiates some earlier reports on the species Imevbore and Bakare, 1970; Imevbore and Okpo, 1975 and Odum, 1992. but conflicts with Adebisi 1988. who described the fish as an invertevore. Variations in the feeding of the species in different habitats indicate flexibility and the accompanying ability to effectively utilize different available food. The broad food spectrum of the species shows that it fed both in the water column and on or near the substratum Table 3.. However C. auratus, like other Chrysichthys species Lowe-McConnell, 1975; Ikusemiju and Olaniyan, 1975., is basically a bottom feeder. This is shown by the significant contribution of dipteran and trichopteran larvae, shrimps and molluscs in the diet. The sandgrains which contributed substantially to the gut contents might have

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been picked up along with food. Their contribution to the nutrition of the species is not clear, but bacteria and protozoans usually associated with sandgrains may be of nutritional benefit. Other food items such as copepods, algae and molluscs were mainly supplementary. All size groups of C. auratus fed on insect larvae with other food items as supplements depending on the size group Fig. 6.. The aquatic insects which are of great importance to all size groups of C. auratus in the River Benin were a secondary source of food for this species in Lake Chad Lauzanne, 1972., where its major food was fish. Fish are known to change their food preference with age Welcomme, 1979. and variations in food habit with size in this study may have resulted from changes in the gape size and food preference of the fish. The temporal food habits show very little qualitative variation but quantitatively there was a distinct variation, more fish being taken during the wet season. This season coincided with the peak breeding period of Tilapia and Brycinus, thus the juveniles of these species form valuable food for C. auratus. C. auratus feed more actively during the night that in the day, an attribute which may account for its increased abundance in night catch as it seems reasonable to assume that most fish are caught when active. Little information on the reproductive biology is available. C. auratus spawned throughout the study period. However, peak spawning occurred in July during the rising flood. The flood among other things provides expanded habitat and abundant food resources to ensure the survival of the huge number of young fishes produced. C. auratus appears to be a one-time spawner in any breeding season, as shown by the uniform state of the eggs in a matured ovary. The fecundity of C. auratus appears low when compared with C. walkeri which at sexual maturity has a size range of 13.324.5 cm total length and fecundity range of 8964168 Ikusemiju, 1976.. Fecundity of C. auratus fluctuated greatly with the body weight and length of fish. Fecundity is known to fluctuate widely even among fish of the same species and size range Bagenal, 1957., and the fluctuations observed in this study may be due to differential feeding success of members of the population prior to spawning.

Acknowledgements We are grateful to Mr. C.B. Powell for confirming the identification of Chrysichthys auratus and Mrs. Esemieburu Sagay for assistance in the field. We are also thankful to the Delta State University for supporting this research programme.

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