International Journal of Fisheries and Aquatic Studies 2019; 7(3): 267-274

E-ISSN: 2347-5129
P-ISSN: 2394-0506
(ICV-Poland) Impact Value: 5.62 Aspects of the biology of juvenile Aba,
(GIF) Impact Factor: 0.549
IJFAS 2019; 7(3): 267-274 Gymnarchus niloticus (Curvier 1829) from Epe
© 2019 IJFAS
www.fisheriesjournal.com Lagoon, Lagos, Nigeria
Received: 09-03-2019
Accepted: 13-04-2019
FV Oluwale, Ugwumba AAA and OA Ugwumba
FV Oluwale
Hydrobiology and Fisheries
Unit, Department of Zoology, Abstract
University of Ibadan, Ibadan, Gymnarchus niloticus is a commercial and socio-cultural important fish in Nigeria. Nevertheless, its
Nigeria fishery is largely based on the seed catch from the wild. Hitherto, diet preference, feeding strategy,
relative gut length, length – weight relationship and Condition Factor (K) of G. niloticus juveniles were
Ugwumba AAA investigated. Fish samples (n = 222) were collected monthly between June and November 2017 from
Hydrobiology and Fisheries artisanal fishermen at Epe Lagoon. Stomach content analysis revealed that juvenile G. niloticus are
Unit, Department of Zoology, carnivorous on fish, insects and crustaceans. Juveniles exhibited specialist feeding strategy on insects.
University of Ibadan, Ibadan, Overall relative gut length was 0.65 (<1) re-enforcing carnivorous feeding. Total length and body weight
Nigeria of juveniles were extremely significantly correlated, r = 0.52 (p< 0.001), and showed isometric growth.
Mean K was <1 during the study. The predilection for insects by juveniles suggested suitability of insect
OA Ugwumba
protein during culture. Poor condition factor indicated vulnerability of the species; hence the need for its
Hydrobiology and Fisheries
Unit, Department of Zoology,
conservation.
University of Ibadan, Ibadan,
Nigeria Keywords: carnivorous, epe lagoon, feeding strategy, juvenile stanzas, gut morphology

1. Introduction
Ample knowledge of the biology on vulnerable early life stages of fishes is central to
recruitment and productivity of later stages. Deficient data on the biology of early life stages of
fish species especially larval and juveniles may be associated with peak abundance and high
mortality rates attributed to these stages in natural population [1, 2]. Consequently, there is a dire
need to elucidate the driving factors of pre-juvenile and juvenile mortality in the wild.
Adequate data on the biology of these vulnerable early life stages remain indispensable
towards comprehending such underlying factors. This may also furnish information on feeding
habits besides associated feeding structural traits, for instance gut morphomerty, that are basal
requirement for fish survival [3, 4].
Dietary composition and feeding habits of fish often vary with fish species, size, time, season,
habitats and other factors all intrinsic links with the environment. At pre-juvenile and juvenile
stages, dietary preference is one of the most important factors in the assessment of feeding
conditions and their probability of meeting food requirements [5]. To this extent, dietary items
of fish are often determined indirectly based on analysis of stomach contents. There is limited
information on the diets of fish juveniles in the wild [6, 7, 8]. Such information could form the
basis of formulation of artificial diets for such species in culture enclosures. Adequate
knowledge on feeding habits of fish species is imperative for culture, rearing and larval control
[9]
.
Relative gut lengths of fish species are increasingly used in conjunction with stomach content
analysis for robust data on feeding habits of fishes. It is one of the useful indices for inferring
feeding habits of fish species into herbivorous, carnivorous, omnivorous, herbi-omnivorous or
carni-omnivorous based on proportion of the gut length with respect to the total length of fish.
It has found widespread application in fisheries biology [10, 11] although this trait often changes
Correspondence with life stages during development [11, 12].
FV Oluwale The Aba Knife Fish, Gymnarchus niloticus is endemic to tropical Africa fresh waters [13].
Hydrobiology and Fisheries Other common names include Freshwater Rat-tail, Nile Knife Fish, Aba Aba, Aba Knife Fish,
Unit, Department of Zoology,
Abba, Frank Fish and Trunkfish.
University of Ibadan, Ibadan,
Nigeria
~ 267 ~
International Journal of Fisheries and Aquatic Studies

It is a peculiar fish devoid of anal and pelvic fin but with a length weight supplied the answer to the third question.
modified caudal fin resembling that of a rat [14, 15]. Amundsen23 plot revealed the feeding strategies interpreted as
Furthermore, the fin that is involved in locomotion is located either generalist or specialist to answer the fourth question.
on the dorsal side instead of the ventral side like in other The fifth question was addressed based on overall calculated
Knife Fishes (Papyroranus afer, Xenomystus nigri). It belongs average condition factor of the species besides its variations
to Family Gymnarchidae, a mono-specific genus. The fish is with time (months).
not only an economic important fish food but also of great
socio-cultural import in Nigeria [14, 16, 17, 18, 19] which endears it 2. Materials and Methods
as one of the most highly valued freshwater fishes in Nigeria. 2.1 Study area
Despite its aquaculture potentials including rapid growth, high Epe lagoon (Figure 1) lies between longitude N 06° 33.710’
premium, tasteful, seasonal availability of wild growers [20], E 004° 03’.710’ and latitudes N 06° 31.893’ E 003° 31.912’.
the supply of Gymnarchus niloticus relies greatly on wild The lagoon is sandwiched between two other lagoons, the
collection which is insufficient for its demands [21]. Tobor Lekki Lagoon (freshwater) in the east and Lagos Lagoon
22
classified the species as endangered due to the (brackish water) in the west. It has a surface area of 243 km2
indiscriminate collection of both young and parent together. and a maximum depth of 6m. The lagoon opens into the Gulf
Wild collection of the juveniles is also on the increase of Guinea (the sea) via the Lagos Harbor, and the vegetation
because of their dark beautiful body colour coupled with their surrounding the lagoon is of the mangrove swampy type. Epe
ability to swim in both forward and backward directions Lagoon supports a predominantly artisanal fishery activity in
makes them ideal ornamental fish. Lagos and Ogun States in Nigeria.
Most studies on Gymnarchus niloticus have been
concentrated on the adult. Therefore, this study aimed to 2.2 Sample collections
report aspects of the biology of the juvenile of a vitally Samples of Gymnarchus niloticus juvenile (Figure 2) were
important commercial fish G. niloticus inhabiting Epe collected monthly for six months between June and
Lagoon. Five major questions were addressed on its biology: November, 2017 from local fishermen at Chief market fish
(i) what are the primary diets of the juvenile of G. niloticus in landing sites located at Epe Lagoon. The fishermen used cast
Epe Lagoon? (ii) what is the relationship between gut and nets and set net to catch fish samples. Samples were preserved
body lengths of juvenile G. niloticus in Epe Lagoon? (iii) in ice-chest to prevent post- capture digestion, then
what is the relationship between its body length and weight? transported into the laboratory and preserved by deep freezing
(iv) what is the feeding strategist of juvenile G. niloticus in (Temperature – 4oC).
Epe Lagoon? What is the condition factor of juvenile G.
niloticus in Epe Lagoon? To address the first questions, 2.3 Data collection
stomach contents were identified and analyzed using primary Specimens were sorted and identified according to Idodo-
indices (Number, frequency of occurrence and volumetric Umeh24 to remove juveniles of other species erroneously
analysis expressed in percentages; validated by a compound collected with the sample. Total lengths of identified
index, Relative Importance Index. The second question was individual were measured with a ruler (Precision 0.1 mm) and
answered by empirical values of relative gut length grounded weighed using a digital weighing balance (Precision 0.1 g).
on ratio of gut length to body length. Regression analysis of

Fig 1: Epe Lagoon

~ 268 ~
International Journal of Fisheries and Aquatic Studies

Fig 2: Gymnarchus niloticus

The length-weight relationship was described by the formula An indirect method based on length and weight measurements
according to LeCren 25W = aLb, where W = Total body wet were used to determine the condition factor of each individual
weight, L = Total body length, “a” is a constant and “b” = fish according to Anderson and Gutreuter [31]:
allometric factor. This equation was logarithmic transformed
to Log W = Log a + b Log L; this approximate a straight line Weight (g) x 105
Condition factor =
when plotted on a graph [26]. Stomach contents were examined [Length (mm)] 3
after internal dissection of the gut; and analyzed based on the
number, volume and frequency of occurrence of each dietary Where w =gutted somatic weight and L = total length
component identified expressed as percentages [27]. Relative
Importance Index (RI) was calculated for each prey and group 2.4 Statistical analysis
of preys based on the Absolute Importance Index (AI) [28]. Gut Descriptive statistics (Mean, standard deviation) was used to
lengths of specimen were measured with a rule (precision 0.1 summarize morph metric parameter of fish data using
mm) after internal dissection. The Relative Gut Length (RGL) Microsoft excel. Amundsen plots of feeding strategy was
was expressed as the ratio of total length of gut to total body plotted using Microsoft excel spreadsheet coded with Macro
length [29]: command 2007. Correlation and linear regression were
performed using Graph Pad Prism Version 5.00 for Windows,
Total length of gut Graph Pad Software, San Diego California USA,
RGL =
Total length of fish www.graphpad.com.

Where fish can be classified as herbivorous (RLG>1), 3. Results

carnivorous (RLG<1) or omnivorous (RLG =, or>, or <1). Body size ranges
The feeding strategy was analyzed according to graphical Total length of juveniles of Gymnarchus niloticus (n = 222)
method of Amundsen [23]; a modification of Costello [30] collected from Epe Lagoon during the sample period ranged
method. This is based on a two-dimensional depiction, in from 49 – 72 mm total length (M = 61.48, SD = 3.59);
which each dietary item point pooled as taxonomic category weighed 0.9 - 0.31g (M 0.52, SD = 0.11).
is generated by plotting the frequency of occurrence (In
fractions) against prey-specific abundance (Pi). Prey specific Dietary composition
abundance is an index of measure of food item based on the In the 222 stomachs examined, nearly half (49 %) had empty
bulk (Volume). Pi was calculated using the volume of food stomachs. Insect parts were the major food of the juveniles of
categories as follows: Gymnarchus niloticus (Table 1); it accounted for 24 %, 18 %
and 0.6 % by number, occurrence and volume respectively.
∑Si x 100 Fish parts were the next importance food in the diet,
Pi =
∑St accounting for 18 %, 4 % and 0.4 % by number, occurrence
and volume respectively. All based on traditional indices
Where: Pi = prey specific abundance, Si = volume of prey i in (Table 1). Based on the relative importance index (Figure 3),
stomachs, and St = total volume of stomach contents in only the primary foods of G. niloticus juvenile are fish, insect and
those individuals with prey i in the stomachs. In order to decapods crustaceans. The most important food category in
determine the feeding strategy (specialized or generalized), the diet of G. niloticus also based on the relative important
dietary items encounter in the stomach were grouped as fish, index (Figure 4) was insects (36.23 %); followed by fish
crustaceans, insect, protozoan and rotifera. (31.96%), while rotifera was the least (6.27 %).

Table 1: Summary of stomach contents of Gymnarchus niloticus juvenile in Epe Lagoon

% Number % Occurrence % Volume Absolute Importance
Food items
(%N) Frequency (%FO) (%V) Index = %N+%FO+%V
Rotifera
Trochocera 5.9 5.3 0.19 11.4
Synchaeta 3.4 3.9 0.09 7.4
Protozoans
Paramecium 2.5 1.3 0.07 3.9
Volvox 7.6 7.9 0.19 15.7
Polytoma 4.2 6.6 1.40 12.2

~ 269 ~
International Journal of Fisheries and Aquatic Studies

Didinium 3.4 2.6 0.47 6.5

Insecta
Insect pupa 5.1 7.9 21.03 34.0
Chironomid larva 3.4 3.9 9.35 16.7
Insect parts 20.3 23.7 14.02 58.0
Fish
Fish parts 16.9 21.1 28.04 66.0
Fish scales 15.3 5.3 9.35 29.9
Crustacean
Copepod 11.9 10.5 15.89 38.3

Fig 3: Diet preference of juvenile Gymnarchurs niloticus based on relative importance index

Fig 4: Relative important indices of food categories in the stomach of juvenile Gymnarchus niloticus in Epe Lagoon

Feeding strategy
Amundsen’s plot (Figure 5) revealed that Gymnarchus Length –weight relationship
niloticus juvenile utilize a specialist feeding strategy. Total length and body weight of juvenile Gynarchus niloticus
Protozoans and rotifers are located in the lower corners were extremely significantly correlated, r (221) = 0.52, p<
indicating abject food items both with low frequencies of 0.001). The results of simple linear regression indicated that
occurrence and prey specific abundance. Juvenile of weight could be predicted from height by the following
Gymnarchus niloticus juvenile is specialist feeders on insects, formula: Y = 3.109x – 3.0476, R2 = 0.2727. The regression
then fish but seldom on crustacean. growth coefficient b = 3.0 shows isometric growth.

Relative gut length

The estimated relative gut length was 0.65.

~ 270 ~
International Journal of Fisheries and Aquatic Studies

Fig 5: Feeding strategy of juvenile Gymnarchus niloticus in Epe Lagoon

Fig 6: Length-weight relationship of juvenile Gymnarchus niloticus from Epe Lagoon

Condition factor
The mean condition factor for the collected juvenile sample of Gymnarchus niloticus was 0.23 ± 0.045. Condition factor of
juveniles varied with time; monthly mean condition factor was generally less than 1 during the present study. Lowest condition
factor was in August 2018 (0.21).

Fig 7: Monthly changes in mean condition factor of juvenile Gymnarchus niloticus from Epe Lagoon

4. Discussion larva is a secondary food for this stage of G. niloticus in Epe

The stomach contents of juvenile Gymnarchus niloticus Lagoon. Oladosu and Oladosu [32] reported that the pre-
comprised five food categories namely fish, insect, rotifer, juvenile feed on the larvae of Chironomid; which is in line
protozoan and crustaceans. This is in agreement with the with observation in this study that chironomid larvae are
reports by other workers [23, 32] that G. niloticus juvenile feed secondary food for juvenile G. niloticus. The midge lay their
on insects while the sub-adults feed on a mixed diet of insects eggs on the grasses that the parent G. niloticus gather around
and fry of other fish. Through the present study, Chironomous their nest [33], and the eggs hatch in a few days. Chironomids
~ 271 ~
International Journal of Fisheries and Aquatic Studies

larvae feed normally on organic matter at the bottom of water temperate and tropical areas [47, 48, 49]. Furthermore,
bodies [34] and are found attached to the submerged grasses indiscriminate dredging and sand filling activities in the
around the nest from where the G, niloticus larvae and lagoon habitats had likely adversely alter bottom condition
juvenile feed on them. Gymnarchus niloticus breeds during over sizeable areas. These could impact negatively on habitat
the flood period [35]; rainy season in Nigeria. The flood period structure and feeding of sensitive and vulnerable juveniles of
is often associated with release of nutrients, rapid growth of this species in the lagoon.
vegetation, and consequently, an elevated availability of other
food sources such as insects, molluscs, seeds, young shoot, 5. Conclusion
leaves [36]. Poor condition of the juveniles indicated vulnerability of the
Examination of distribution of points along the diagonal and species, and hence the need for its conservation in Epe
axes the diagram (Figure 6) of the Amundsen’s plots showed Lagoon. Furthermore, the prevailing condition of Epe Lagoon
clearly a specialist feeding strategy for juvenile G. niloticus in may be unsuitable for the juveniles as indicated by pollution
Epe Lagoon. This specialist feeding strategy comprises two indicator, chironomid, encountered in their stomachs as food
components: constant or primary food (insect, fish and items. Insectivorous habits of the juveniles implied that the
crustacean) and occasional food (rotifers and protozoan). The species may thrive on insect protein diet during culture.
dietary preferences in fish rely mainly depend on the nature of
food available in the living habitat, environmental conditions, 6. References
size or sexual stages of fish as well as inter and intra specific 1. Rothschild B. Dynamics of marine fish populations.
competition [37, 38]. Harvard University Press, Cambridge, 1986, 97.
Juvenile stages of Gymnarchus niloticus are carnivorous as 2. Houde. Fish early life dynamics and recruitments
indicated by a Relative Gut Length (RGL) index of 0.6. This variability. American Fishery Society Symposium. 1987;
is not surprising since diet of the stage consists of strictly 2:17-19.
materials of animal origins as observed in the present study. 3. Balon EK. Terminology of intervals in fish development.
In general, gut length varies predictable with diet: carnivores Journal of the Fisheries Research Board of Canada. 1975;
< omnivores < herbivores. This assertion is based on the 32:1663-1670.
ratios of alimentary tract length to body length for fishes in 4. Webb PW. Functional locomotor morphology of early
diverse dietary categories. Lagler [39] ascertained that shorten life history Stages of fishes Transactions of the American
intestine in carnivores compare to other feeding group could Fisheries Society. 1986; 115:115-127.
be attributed to ease of digestion of food from animal origin 5. Robichaud-LeBlanc KA, Courtenay SC, Hanson JM.
than vegetable ones. Kramer and Bryant [40] categorized small Ontogenic diet shifts in age of striped bass, Morone
fishes (50–100 mm SL) by RGL as carnivores (RGL=0.6– saxatilis, from the Miramichi River estuary, Gulf of St.
0.8), omnivores (0.8–1.0) or as herbivores (2.5–16.4). With Lawrence. Canadian Journal of Zoology. 1997; 75:1300-
respect to feeding, the alimentary tract of fishes displays 1309.
anatomical and physiological adaptation 41, 42 also reflected in 6. Humphries P, King AJ, Koehn J. Fish, flows and
the gut length. floodplains: Links between freshwater fish and their
The result shows that the length of the fish increased with environment in the Murray-Darling Rive system,
body weight. Gymnarchus niloticus juvenile exhibit isometric Australia. Environmental Biology of Fishes. 1999;
growth (b=3.0). Isometric growth implies all body parts grow 56:129-151.
at almost the same rate as the fish increases in size. Extremely 7. Kakareko T, Zbikowski J, Zytkowicz J. Diet partitioning
significant correlation between body length and body weight in summer of two syntopic neogobiids from two
also support this assertion. Wootton [43] argued that fish differenthabitats of the lower Vistula River, Poland.
exhibiting isometric growth pattern will follow the cube law; Journal of Applied Ichthyology. 2005; 21:292-295.
retain body shape with constant specific gravity. Hence, b- 8. Ara R, Arshad A, Musa L, Amin SMN, Kuppan P.
value must be equal to 3.0. In contrast, negative allometric Feeding habits of larval fishes of the family Clupeidae
growth (b<3) [44, 45] were reported for adult G. niloticus. This (Actinopterygii: Clupeiformes) in the Estuary of River
indicates as fish growths it becomes lighter for its length. Pendas, Johor, Malaysia. Journal of Fishery and Aquatic
Differences in growth patter b for juvenile and adult’s G. Sceinces. 2011; 56:816-821.
niloticus is of course expected. These differences indicate that 9. Ndome CB, Victor R. Food and feeding habits of
the species is passing through stages in its life cycle defined Epiplatys senegalensis (Pisces: cyprinodontiformes;
by different length – weight relationship. This agreed with Cyprinodontidae) in a back water pond in Benin City,
Ayoade [44] who reported that the different growth pattern was Southern Nigeria. West African Journal of Applied
demonstrated by the size groups of Schilbe mystus in Asejire Ecology. 2002; 3(1):105-117.
Lake. Juvenile and adult stages of a fish may exhibit 10. Dasgupta M. Morphometrics of the alimentary canal of
differences in the length – weight relationships owing to the some freshwater fishes of West Bengal in relation to their
changes in the body form with size, feeding habits and factors food and feeding habits. Indian Journal of Fisheries.
related to reproduction [46]. 2002; 49(4):461-464.
There was temporary variation in condition factor of juvenile 11. Koundal S, Dhanze R, Koundal A, Sharma I. Relative gut
Gymnarchus niloticus inhabiting Epe Lagoon. The fact that length and gastro-somatic index of six hill stream fishes,
the average condition factor was <1 suggested during the Himachal Pradesh, India. Journal of Environment and
present study indicates that juvenile stage G. niloticus might Biosciences. 2013; 27(1):11-18.
be in poor condition at Epe Lagoon. A pollution indicator, 12. Bhuiyan AS, Islam MN. Observation on the Food and
Chironomus larva, was also a food item indicating possible Feeding Habit of Ompok pabda (Ham.) from the River
pollution in the lagoon. High abundance of chironomids is a Padma. Pakistan Journal of Zoology. 1991; 23:75-77.
confirmed phenomenon in polluted water bodies in both 13. Nelson JS, Grand TC, Wilson MVH. Fishes of the World.

~ 272 ~
International Journal of Fisheries and Aquatic Studies

5 ed. Hoboken, New Jerssy John Willey and Sons Inc, 32. Oladosu GA, Oladosu OO. Investigating suitable first
2016, 752p. food sources for hatchery rearing of Gymnarchus
14. Reed W, Barchard J, Hopson AJ, Jennes J, Yaro I. Fish niloticus (Cuvier 1829) fry. Journal of Agriculture
and fisheries of Northern Nigeria. Edition 1, Ministry of Forestry' and Fisheries. 2002; 3:13-14.
Agriculture, Northern Nigeria, 1967, 31-32. 33. Oladosu GA. Environmental induction of 'natural
15. Paugy D, Lévêque C, Teugels GG. The Fresh and spawning in Gymnarchus niloticus (Cuvier 1829) in an
Brackish Water Fishes of West Africa. (1). Institut de earthen pond. Aquaculture Research. 1997; 28:641-643.
recherché pour le developpement Paris France. Musée 34. Burgis MJ, Morris P. The natural history of lakes.
royal de l'Afrique Centrale Tervuren, Belgigue, 2003, Cambridge University Press, 1987, 212.
457p. 35. Bennett MVL. Electric Organs. In: Fish Physiology,
16. Sagua VO. Preliminary reports on the biology of Hoar, W.S. and D.J. Randall (Eds.). 9th Edn. Academic
Gymnarchus niloticus in Lake Chad. In: Annual report of Press, London, 1971, 347-491.
Lake Chad Research Institute, Nigeria, 1983, 61-65. 36. FAO/UN. Report to the Government of Nigerian on
17. Oladosu GA. Environmental induction of 'natural fishery investigations on the Niger and Benue rivers in
spawning in Gymnarchus niloticus (Cuvier 1829) in an the Northern Region and development of a program of
earthen pond. Aquaculture Research. 1997; 28:641-643. riverine fishery management training. Based on the work
18. Oladosu OO, Oladosu GA, Hart AI. Some ecological of M.P. Motwari Rep. FAO/UNDP(TA), (2771), 1970,
factors of the tropical floodplain on the breeding and 196P.
conservation of Gymnarchus niloticus. In Kolo, R.J. and 37. Zacharia PU, Abdurahiman KP. Methods of stomach
Orire A. M. (eds). Proceedings of the 26th Annual content analysis of fishes. CMFRI, 2004, 148-158.
conference of the Fisheries Society of Nigeria. Minna, 38. Lawal MO, Edokpayi CA, Osibona AO. Food and
2011, 193-200. feeding habits of the guppy, Poecilia reticulata, from
19. Ayoola SO, Abotti CE. Morphology of the amba knife drainage canal systems in Lagos, Southwestern Nigeria.
fish (Gymnarchurs niloticus (Curvier, 1829) World West African Journal of Applied Ecology. 2012; 20(2):1-
Journal of Fisheries and Marine Science. 2010; 2(5):354- 9.
356 39. Lagler KF, Bardach JE, Miller RR, Passino DRM.
20. Kigbu AA, Imgbian TD, Yakub MM. Unconventional Ichthyology. 2nd Edn. John Wiley and Sons, New York,
cultivable freshwater fish species: a potential tool for 1977, 129-170.
increased production in Nigeria Global Science Research 40. Kramer DL, Bryant MJ Intestine length in the fishes of a
Journal. 2014; 2(3):152-157. tropical stream: 1. Ontogenetic allometry. Environmental
21. Andrew G, Boyd AH, Michael MJV. Aquaculture Biology of Fishes. 1995; 42:115-127.
development and research in the sub-saharan Africa. 41. Cortés E, Papastamatiou YP, Carlson J, Ferry LA,
CIFA Tech. Paper, 1994, 23. Wetherbee BM. An overview of the feeding ecology and
22. Tobor JG. Fin and shellfish of conservation interest in physiology of elasmobranch fishes. In: Feeding and
Nigeria. Nlotvlk Tech. Paper No. 1992, 79. digestive functions of fishes. (eds J.E.P. Cyrino, D.P.
23. Amundsen PA, Gabler HM, Staldvik FJ. A new approach Bureau, B.G. Kapour) Science Publishers, Enfield, New
to graphical analysis of feeding strategy from stomach Hampshire, 2008, 575p.
contents data-modification of the Costello (1990) 42. Yashpal M, Kumari U, Mittal S, Mittal AK.
method. Journal of Fishery Biology. 1996; 48:607-614. Morphological specialization of the buccal cavity in
24. Idodo-Umeh, G. Freshwater Fishes of Nigeria. Idodo- relation to the food and feeding habit of a carp Cirrhinus
Umeh Publishers Limited Benin City, 2003, 231. mrigala: a scanning electron microscopic investigation.
25. LeCren ED. The length-weight relationship and seasonal Journal of Morphology. 2009; 270(6):714-728.
cycle in gonad weight and condition in the Perch (Perca 43. Wootton RJ. Ecology of Teleost Fishes. 1st Edn.,
fluviatilis). Journal of Animal Ecology. 1951; 20:201-219 Chapman and Hall, London, UK., ISBN-13:
26. Weatherly AH. Growth and ecology of fish populations. 9780412317200, 1990, 404.
Academic Press, London, 1972, 72. 44. Ayoade AA. The length – weight relationship of Schilbe
27. Hyslop EJ. Stomach contents analysis- a review of mystus from (Linne, 1766) two man-made lakes in south-
methods and their applications. Journal of Fish Biology. western Nigeria. The Zoologist. 2011; 9:38-43.
1980; 17:411-429. 45. Odo GE, Dibua E, Ekeh FN, Ivoke N, Asogwa CN,
28. George EL, Hadely WH. Food and habitat partitioning Avoaja DA, Atama. The breeding biology and
between rock bass (Ambloplites rupestris) and small hematological profile of Gymnarchus niloticus Cuvier
mouth bass (Micropterus dolomieu) young of year. 1829 (Osteoglossiformes: Gymnarchidae) in semi-
Transactions of American Fish Society. 1979; 108:253- intensively managed ponds in the floodplain of the River
261. Anambra, Nigeria. Journal of Fisheries and Aquaculture.
29. Biswas SP. Manual of methods in fish biology. The South 2013; 4(2):103-109.
Asian publishers Pty ltd. 3 Nejati subhoshmary. 46. Khan Afzal, Khan MS, Miyan K. Length – Weight
Daryagam, New Dehli, India, 1993, 157P. relationship of giant snakehead, Channa marulius and
30. Costello MJ. Predator feeding strategy and prey stinging catfish, Heteropneustes fossilis from the River
importance: A new graphical analysis. Journal of Fishery Ganga India. Journal of Applied Ichthyology. 2012;
Biology. 1990; 36:261-263. 28:154-155.
31. Anderson RO, Gutreuter SJ. Length, weight, and 47. Cardoso I, Granadeiro JP, Cabral H. Benthic
associated structural indices. In Nielson LA, Johnson DL, macroinvertebates vertical distribution in the Tagus
editors. Fisheries techniques. Bethesda, Maryland: estuary (Portugal): The influence of tidal cycle. Journal
American Fisheries Society, 1983; 283-300. Estuarine Coastal Shelf Sciences. 2010; 86:580-586.

~ 273 ~
International Journal of Fisheries and Aquatic Studies

48. Edward JB, Ugwumba AAA. Macroinvertebrate fauna of

a tropical southern reservoir, Ekiti State, Nigeria.
Continental Journal of Biological Sciences. 2011;
4(1):30-40.
49. Yakub AS, Ugwumba AAA. A study of the
macroinvertebrate fauna of lower Ogun River at Ishasi,
Ogun State, southwest Nigeria. The Zoologist. 2009;
7:65-74.

~ 274 ~