463

A review of on-farm feed

management practices
for North African catfish
(Clarias gariepinus)
in sub-Saharan Africa

Thomas Hecht1
Department of Ichthyology and Fisheries Science
Rhodes University, Grahamstown
South Africa

Hecht, T. 2013. A review of on-farm feed management practices for North African
catfish (Clarias gariepinus) in sub-Saharan Africa. In M.R. Hasan and M.B. New,
eds. On-farm feeding and feed management in aquaculture. FAO Fisheries and
Aquaculture Technical Paper No. 583. Rome, FAO. pp. 463–479.

Abstract
This review considers feed management practices for North African catfish
(Clarias gariepinus) in sub-Saharan Africa. Clariid catfish production in the
subcontinent is increasing exponentially, particularly in Uganda. Semi-intensive
pond culture is still the most prevalent production system, while intensive tank
culture is becoming more popular in peri-urban areas in Nigeria. Total production
in these two countries now exceeds 178 000 tonnes annually. Catfish are now
commonly spawned and their larvae reared in hatcheries for ten to 14 days, after
which they are reared in nursery ponds or in tanks. Extensive rearing of larvae,
after yolk sac absorption, in ponds is now less often practiced than in the past.
Where it is still practiced, successful larval rearing and satisfactory survival rates
depend mainly on adequate fertilization schedules. Feed management practices
in hatcheries are closely matched with the physiological and endocrinological
ontogeny of the fish. For optimal survival and growth, live food (mainly
Artemia) is required for the first five days after the start of exogenous feeding,
after which the fish can be weaned onto a dry starter feed. Up to a size of
5 g, the species has a high protein demand (>50 percent). Extensive farming
of catfish in ponds is largely a subsistence activity and is practiced mainly in
polyculture with tilapia that serve as fodder fish, and using a single ingredient
feed such as maize or wheat bran. Semi-intensive on-growing of catfish in static
and flow-through ponds, as well as under high-density tank culture conditions
requires a complete feed; production levels achieved in these three systems are
15–24 tonnes/ha/cycle, 40 tonnes/ha/cycle and 385 kg/m3/cycle, respectively.
Results show that floating extruded pellets with a protein content of 30–35 percent
are preferred by farmers. The duration of the grow-out cycle depends on the size
of fish required by the market. At temperatures between 26 and 28 oC the fish can
be grown from 1 g to 800 g in seven months. Feed conversion ratios (FCR) are

1
Present address: Advance Africa Management Services, PO Box 381, Port Alfred. South Africa.
464 On-farm feeding and feed management in aquaculture

size dependent, and best ratios are obtained by feeding the fish to satiation while
observing their feeding response. Daily ration tables serve largely as a guideline.
In ponds, the fish are fed two to three times per day, while under high-density
tank conditions they are fed five to six times per day. During the early juvenile
phases (1–24 g), FCR are commonly <1:1; from 25 g to 800 g FCR of 1.2:1 are
achievable. There have been significant advances in feed availability and quality
in the region, particularly in Uganda, although it would appear that weaning
diets and starter crumbles are still being imported.

1. Introduction
While the aquaculture attributes of the North African or sharptooth catfish, Clarias
gariepinus have been recognized since the 1940s (Hey, 1941), the basic technologies
and protocols for the farming of the species were developed much later, mainly from
the 1960s to the late 1990s (e.g. Micha, 1971, 1972; El Bolock, 1973; De Kimpe and
Micha, 1974; Richter, 1976; Hogendoorn, 1979, 1981; Bok and Jongbloed, 1984; Viveen
et al., 1985; Verreth and Den Bieman, 1987; Hecht, Uys and Britz, 1988; De Graaf and
Janssen, 1996,). These studies provided the basic information upon which the future
research and farming of North African catfish would be founded. Many mistakes were
made along the road to commercialization (Hecht, Oellermann and Verheust, 1996),
and it is only in the more recent past (i.e. in the first decade of the new millennium) that
C. gariepinus farm production figures for sub-Saharan Africa (Table 1) have become
significant (Figure 1), particularly in Nigeria and latterly in Uganda (Figure 2). The
growth of the catfish farming industry in Nigeria was market driven, while the spike in
production in Uganda in 2006 is generally regarded as a combination of market forces
and a well-directed donor project initiative (Isyagi et al., 2009a). According to FAO
(2012), North African catfish was the most important commercial aquaculture species
in 2010 in sub-Saharan Africa (198 296 tonnes), followed by Nile tilapia (Oreochromis
niloticus) (60 350 tonnes). The farming of C. gariepinus in Europe is a minor activity
(FAO, 2012) and is currently occurring only in the Netherlands, Hungary and Poland
(1 810, 3 200 and 650 tonnes, respectively, in 2010). In Asia, C. gariepinus is hybridized
with C. batrachus and C. macrocephalus. Clariids are an important component of
the total farmed fish production in the Asia Region, which in 2010 contributed a
production of 453 209 tonnes (FAO, 2012).

TABLE 1
Total production (tonnes) of clariid catfish in Africa, 2004 to 2010

2004 2005 2006 2007 2008 2009 2010

Egypt 459 10 180 6 058 5 287 13 944 17 895 9 717

Kenya 320 318 302 890 935 1 047 2 188

Nigeria 26 750 34 582 51 916 52 229 86 130 89 193 130 318

Uganda 3 827 6 528 20 941 34 096 35 000 54 956 63 000

Other (n=22) 721 617 922 1 037 1 159 1 669 2 790

Sub-Saharan subtotal 31 618 42 045 74 081 88 252 123 224 146 865 198 296

Total 32 077 52 225 80 139 93 539 137 168 164 760 208 013

Source: FAO (2012).

Feed management practices for North African catfish in sub-Saharan Africa 465

FIGURE 1
Clariid catfish production in Africa, 1990 to 2010

Source: FAO (2012).

FIGURE 2
Percent contribution by major1 sub-Saharan clariid catfish producer countries2

1
Defined as having a production of >1 500 tonnes in 2010.
2
The ‘other’ category consists of 22 sub-Saharan countries in which clariid catfish are also produced in lesser
quantities.

Source: FAO (2012).

466 On-farm feeding and feed management in aquaculture

The significant increases in annual production in sub-Saharan Africa from around

2000 onwards can be partially ascribed to the concerted research efforts on the nutritional
requirements of the species during all life history stages, feed formulation, optimization
of feeding and feed management practices and the development of high-density tank
farming practices. However, more importantly, there has been a switch in the profile of
catfish farmers from small-scale rural farmers to educated, mainly urban-based, young
entrepreneurs who have easier access to capital and for whom access to information is less
difficult. In sub-Saharan Africa, clariid catfish are farmed in one of three basic ways viz.
extensive polyculture in ponds, semi-intensive pond culture and intensive tank culture.
In smallholder ponds, the fish either contribute to some extent towards improved
nutrition at the household level or (when produced in surplus), are (rarely) sold for cash.
The fish are normally produced in polyculture with tilapiine species that serve as fodder
fish. The most common feed used under this scenario is maize or wheat bran (from which
the catfish also benefit marginally), and some (normally inadequate) quantity of manure,
compost and possibly some fertilizer are applied. Under this scenario, production rarely
exceeds 1.5 tonnes/ha/year, which equates to ~30 kg of fish per 200 m2 pond per annum.
Commercial-scale production of catfish in ponds takes place in various ways.
Under monoculture, static pond or flow-through conditions, the fish are normally
fed on a complete, pelleted feed or on farm-made feeds. Production levels under static
pond conditions range from 15–20 tonnes/ha/cycle. Under flow-through conditions,
production ranges between 25 and 40 tonnes/ha/cycle. In polyculture conditions in
stagnant ponds, catfish are often grown together with O. niloticus and fed on farm-
made feeds, with total yields ranging from 5 to 8 tonnes/ha/year (Ayinla, 2007). During
the period 1985 to 1998, there was a concerted research effort on high-density tank
farming of Clarias gariepinus under recirculating conditions (Oellermann and Hecht,
1996; Eding and Kamstra, 2002; Eding et al., 2006). This technology was first put
into practice in the Netherlands and is now commonly used in Nigeria, Hungary and
Poland. It should, however, be noted that the bulk of catfish in Nigeria and Uganda
is still produced in ponds (Akinwole and Faturoti, 2007; J. Rutaisire, Department of
Fisheries, Uganda, personal communication 2010). High-density tank farms in Nigeria,
and now also in Uganda, are situated mainly in peri-urban environments and in other
areas where there is ready access to supplies and services (Akinwole and Faturoti, 2007).
In Uganda and Kenya, C. gariepinus is not only farmed commercially as a food fish,
but fingerlings are also produced as live baitfish for the Nile perch (Lates niloticus)
fishery in Lake Victoria. Currently the demand for 8–10 g fingerlings as live bait is
around 700 million per year in the Lake Victoria basin area, while the demand in
Uganda alone is around 300 million per year (Isyagi et al., 2009a, b).
This paper reviews current and best on-farm feeding and feed management
practices during the various life history stages. Based on current practices, several
recommendations are provided within the text that may help to improve production
and feed conversion ratios (FCR) and lead to reduced levels of waste.

2. FEEDING OF LARVAE AND EARLY JUVENILES

Catfish larvae are generally reared to fingerlings in one of three ways;
• Under commercial farming conditions, they are either intensively reared to
fingerling size solely in a hatchery, after which they are on-grown under pond or
high-density tank culture conditions.
• Alternatively, they are reared in a hatchery for a period of up to 14 days and then
grown to fingerling size for a further 30 + days in nursery ponds, after which they
are stocked into larger on-growing ponds.
• Extensive larval rearing encompasses the stocking of 3-day-old larvae into well
prepared and protected nursery ponds, where they are on-grown to fingerling size
before being stocked into larger grow-out ponds.
Feed management practices for North African catfish in sub-Saharan Africa 467

The feed to be used and the applied feeding strategy are largely dependent on
whether the larvae are reared extensively or intensively.

2.1 Extensive rearing

Under extensive rearing conditions, the ponds are prepared such that there is
an abundance of zooplankton. This takes place up to 14 days prior to stocking
and consists of liming and fertilization. ASEAN (1978) and Viveen et al. (1985)
recommend the use of 100–150 kg/ha of quicklime, which is added to the damp
pond bottom to eliminate pathogens and invertebrate predators. The pond is then
left for 7–14 days, after which it is filled with water to a depth of 30 cm and the pH
of the water adjusted by adding agricultural lime. Ensuring a good zooplankton
bloom requires adequate fertilization with chemical or organic fertilizers. The choice
of either is largely determined by availability and the economic circumstances of
the farmer. The main inorganic fertilizers used are super-phosphate (approximately
20 percent P2O5), triple super-phosphate (45 percent P2O5), urea (about 45 percent N)
and NPK 15:15:15 (15 percent N, 15 percent P2O5 and 15 percent K2O). It is
strongly recommended to dissolve the fertilizers and then to spread the solution
evenly over the water surface (ASEAN, 1978). In the absence of any specific
work on North African catfish fry or fingerling rearing ponds, it is advisable to
follow the findings and recommendation made by Hepher (1963, 1967) and Boyd
(1982). The work by Hepher (1963, 1967) has shown that applying fertilizer doses
greater than 0.5 mg P/L or 1.4 mg N/L has no biological or economic justification.
These levels are equivalent to application rates of 60 kg/ha of single super-phosphate
(11 kg P2O5/ha) and 60 kg/ha ammonium sulphate (13 kg N/ha) respectively, applied
at two-weekly intervals (0.8–1.0 m water depth, 8 000–10 000 m3 water/ha) (Tacon,
1987, 1988). Similarly, Boyd (1982) recommends chemical fertilization strategies to
maintain soluble nitrogen and orthophosphate at 0.95 mg N/L and 0.1–0.5 mg P/L,
respectively.
The most commonly used organic fertilizers are chicken, pig and cow manure.
Table 2 shows several application schedules. It is important to note that the application
rates are not absolute and only serve as guidelines. If an adequate phytoplankton
bloom is not achieved within six to eight sunny days, then more manure should be
added. Plankton levels are adequate when the water has a green colour and a Secchi disk
reading of around 20–25 cm. The biological effect of organic fertilization is improved
if the manure is spread evenly over the water surface in a liquid or semi-liquid form
(Tacon, 1988). Fish ponds can only assimilate a certain amount of manure per day.
Hence to prevent de-oxygenation in manure-loaded ponds (due to unchecked peaks
in bacterial growth and phytoplankton blooms), manure should be added frequently
(daily, if possible) and in small amounts, and during mid-morning when oxygen levels
are rising (Hepher and Pruginin, 1981).
Table 3 shows a mixed (inorganic and organic) fertilization schedule that can also be
used to develop good zooplankton blooms.

TABLE 2
Fertilization rates using animal manure

Fertilizer Rates (kg dry weight/100 m2)

Poultry manure 25 kg initially and then 3–5 kg every 10 days

Pig manure 7 kg every 2nd day

Cow dung 10 kg every 2nd day

Source: Tacon (1988).

468 On-farm feeding and feed management in aquaculture

TABLE 3
Combined fertilization rates
Manure/fertilizer Rate (kg/week/100 m2)
Dry poultry manure 10–20

Urea 0.4–0.8

Triple superphosphate 0.1–0.2

Source: Janssen (1985).

Periphyton can also be used successfully for the rearing of Clarias larvae
(Nwachukwa, 1999). Amisah, Adjei-Boateng and Afianu (2008) have demonstrated
the beneficial combined effect of pond fertilization and the use of bamboo poles for
the development of periphyton. In their study, bamboo pole density was 4 per m2 and
ponds were fertilized with pig manure at an initial rate of 20 kg/100 m2 and then every
second week at 10 kg/100 m2.
Clarias larvae start exogenous feeding approximately 80 hours after hatching
(depending on temperature) and are therefore normally transferred into well prepared
and protected nursery ponds when they are three days old. Nursery ponds vary in
size between 100 and 250 m2; in sub-Saharan Africa nursery ponds must be protected
against the African clawed toad, Xenopus laevis, by way of a smooth wall made of
plastic, roof sheeting or other suitable material. If this precaution is not taken, then
the majority, if not all, of the fry will be lost to predation. The ponds are stocked
at a density of 100–250 larvae/m2. From the day of stocking, the fish must receive
supplementary feed. At a density of 100 fry/m2 the recommended feeding levels are
1 kg rice or wheat bran, plus 1 kg/100 m2 of crumbled, formulated feed containing
some fishmeal for the first three weeks. For the following two weeks, the bran ration
remains the same but the formulated feed ration increases to 2 kg/100 m2/day, fed in
two equal meals. After three weeks, it is advisable to grade the fish into two or three
size groups to reduce cannibalism. With good feeding and management practices,
survival rates of 40 percent and a final fish weight of 3 g can be achieved during 50-day
rearing cycles (De Graaf and Janssen, 1996). These authors are also of the opinion that
the most critical factor (on condition that the pond is protected) is most probably the
availability of zooplankton during the first few days of the rearing cycle.

2.2 Intensive rearing

The larval rearing protocol under intensive hatchery rearing conditions is completely
different and is very closely allied to the ontogeny of the alimentary and endocrine
systems. It normally lasts for a period of 12 to 14 days and ideally takes place at the
optimum temperature for growth (28 °C).
Research has shown that live food in the form of enriched Artemia nauplii, small
Daphnia, Moina or other zooplankton of suitable size is essential for the first four
to six days after the start of exogenous feeding (Hogendoorn, 1980; Uys and Hecht,
1985; Verreth and van Tongeren, 1989; Haylor, 1993; Hecht, 1996; Awaiss and
Kestemont, 1998) and is preferable to dry food only (Appelbaum and van Damme,
1988). The reason for this is that the stomach of the larvae is not functional at the
start of exogenous feeding (three days after hatching), and its development extends
beyond the eleuthero-embryo stage into the larval stage (Stroband and Kroon, 1981).
The stomach only becomes functional four to five days after the start of exogenous
feeding (Verreth et al., 1992; Segner et al., 1993). Once the stomach becomes
functional and pepsin activity contributes significantly to protein digestion, the
larvae can be weaned from live food to a dry feed (Verreth and van Tongeren, 1989;
Verreth et al., 1993).
Feed management practices for North African catfish in sub-Saharan Africa 469

Larvae and early juveniles of C. gariepinus (up to approximately 5 g) have a high

protein demand of around 50–55 percent and a lipid requirement of 9 percent (Uys
and Hecht, 1985), of which at least 10 percent should consist of fish oil (Uys, 1989;
Kerdchuen, 1992). The carbohydrate content can be as high as 21 percent of the diet
(Uys and Hecht, 1985). For optimal growth, the composition of any dry feed that will
be fed from 5 or 6 days after hatching (dah) until at least 14 dah must meet the nutrient
requirements as outlined above.
Several larval and early juvenile rearing protocols have been developed. The four
most commonly used protocols are shown in Table 4; all result in good growth and
survival rates. Optimum pellet crumble size during this period is 2.2 percent of the
mean total length of the larvae, and early juveniles should be fed at 25 percent of body
weight per day at two-hour intervals (Uys, 1984). Predicted feeding rates are also
provided by Verreth and Den Bieman (1987). Production costs can be significantly
reduced by using small-strain Daphnia instead of Artemia.

TABLE 4
Four feeding protocols for North African catfish, Clarias gariepinus1
Protocol 1 (dah)2,3 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 15/16–30/32
Artemia

Weaning diet

Grow-out feed

Protocol 2 (dah)4 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 15/16–30/32

Artemia

Weaning diet

Grow-out feed

Protocol 3 (dah)5 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 15/16–30/32

Artemia

Weaning diet

Grow-out feed

Protocol 4 (dah)6 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 15/16–30/32

Artemia

Weaning diet/bran

1
Larvae start exogenous feeding ca. 80 hours after hatching
2
Protocol 1 (Hecht, Uys and Britz, 1988): Artemia is presented to the fish once a day
3
dah = days after hatching
4
Protocol 2 (Verreth, Storch and Segner, 1987): Artemia restricted up to four times per day
5
Protocol 3 (Oellerman, 1995): Artemia restricted up to four times per day
6
Protocol 4 (Janssen, 1985; De Graaf and Janssen, 1996): Artemia presented ad libitum

After 12–14 days, the fry are stocked into nursery ponds at densities ranging from
65 to 2 000/m2 (Viveen et al., 1985; Hecht, Uys and Britz, 1988; Isyagi et al., 2009b).
The density at which fry are stocked depends on fish age and the intensity of rearing
protocols. Under pond-farming conditions, it is recommended to feed the fry three
times per day at 25 percent of body weight per day, using a 38–40 percent protein
diet (Hecht, Uys and Britz, 1988). If the larvae and early juveniles are reared in tanks,
then the feed should have a protein content of around 50 percent. When the fish
attain an average weight of 1–2 g, they are ready to be stocked into ponds or tanks for
on-growing.
470 On-farm feeding and feed management in aquaculture

Cannibalism is a serious problem during larval and early juvenile rearing,

particularly under hatchery conditions (Hecht and Appelbaum, 1988). The rate of
cannibalism is affected inter alia by feeding frequency, food availability, light intensity
and photoperiod (Hecht and Pienaar, 1993). Almazán, Schrama and Verreth (2004)
showed that a reduction in swimming activity results in a lower incidence of aggressive
behaviour. Hence feeding frequency, food availability and light intensity must be
manipulated such that swimming activity is reduced to a minimum. This is achieved
at low light intensity (<15 lux), continuous food availability and feeding the fish every
two hours around the clock.

3. FEEDING DURING ON-GROWING

The species is produced in widely different farming systems ranging from highly
intensive tank culture under flow-through or recirculating conditions to intensive,
semi-intensive and extensive (small and large-scale, polyculture or monoculture)
static or flow-through pond-culture systems. It is therefore not surprising that a wide
range of feeds are used in the farming of this species. These include dry feeds ranging
from single ingredient dry feeds, such as maize bran, to farm-made mixed feeds to
formulated, floating or slow-sinking, extruded pellets, as well as single ingredient or
mixed moist, farm-made feeds (Hecht, 2013).
The optimum pond size for catfish production was found to be 0.1 ha. This allows
for optimum control of feed use and management of the production and harvesting
cycle (Hecht, Uys and Britz, 1988). Under pond-farming conditions using formulated
feeds, it is best to train the fish to feed in the same area of the pond at the same time
of day. This is easily achieved and has several advantages. It allows the farmer to take
good samples with which to assess average fish weight and fish health and ultimately
facilitates harvesting. It is further recommended to feed the fish based on their feeding
response. This allows the fish to be fed to satiation on a daily basis under variable
environmental conditions (e.g. temperature, dissolved oxygen, pH, nitrogenous
metabolites, barometric pressure), all of which affect daily feed intake. Feeding charts
should serve as a guide to estimate daily rations (Table 5). During the on-growing cycle
in ponds, fish are normally fed twice a day. It is an essential management practice to
maintain good records of the daily amounts fed and the feeding response of the fish
(Hecht, Uys and Britz, 1988; Isyagi et al., 2009b).

TABLE 5
Recommended daily ration for North African catfish (expressed as % body weight/day)
Temperature Fish size (g)
(°C)
1–10 10–25 25–50 50–100 100–300 300–800
16 1.0 0.6 0.4 0.3 0.2 0.2

18 3.0 1.6 1.0 0.8 0.6 0.5

20 5.0 3.0 2.0 1.5 1.2 1.0

22 6.8 4.5 3.0 2.4 2.0 1.7

24 8.1 6.0 4.0 3.0 2.5 2.2

26 9.5 6.6 5.1 3.6 3.2 2.8

28 10.0 7.0 5.5 4.0 3.5 3.1

30 9.8 6.8 5.3 3.7 3.2 2.9

32 9.5 6.5 5.0 3.5 3.0 2.8

Source: Hogendoorn et al. (1983); Hecht, Uys and Britz (1988).

Feed management practices for North African catfish in sub-Saharan Africa 471

Under pond-farming conditions, it is advisable to feed the fish at the same time and
at the same place each day such that they become accustomed to a particular feeding
regime. This allows the farmer to feed on a response basis, which results in less waste
and improved FCR (Isyagi et al., 2009b). Under good pond management practices,
FCR during grow-out of 1:1 are not uncommon, although this of course depends on
the nutrient profile of the feed and the aptitude of the farmer.

3.1 Nutrient requirements and feed production and availability

The nutrient requirements of Clarias gariepinus are outlined in Table 6; based on this
information various least-cost feeds of variable quality are available on the market
throughout the region. In the two largest producer countries (Nigeria and Uganda),
catfish feeds are readily available. In Nigeria, there are at least 12 commercial aquafeed
producers, and products range from slow-sinking pellets to floating extruded pellets.
In Uganda, there is one large producer (Ugachick) that produces a slow-sinking as well
as an extruded floating catfish feed (30 percent protein pellets) and, as in Nigeria, there
are many small-scale feed producers. All of the commercial feed producers in Nigeria
essentially target the needs of catfish farmers. However, specifically formulated and
extruded pellets (45 percent protein) for the early rearing stages of C. gariepinus are
imported from South Africa and elsewhere (Ayinla, 2007).

TABLE 6
Dietary nutrient requirements of North African catfish (10–1 000 g)
Nutrient Quantity1 References2
Crude protein (% min) 40–43 1, 2, 3, 4, 5, 6, 7, 8, 9

Least cost/appetite feeding protein requirement 35–38 1, 6

Crude lipid (% min) 10–12 1, 2, 6, 7

Carbohydrate (% recommended) 15–35 1, 6, 7, 10, 11, 12

Digestible energy (min, kJ/g) 14–16 1, 2, 13

Metabolizable energy (min, kJ/g 13 2

Gross energy (min, kJ/g) 22–24 2, 4, 7, 10

Protein to energy ratio (mg/kJ) 22–30 1, 2, 4, 7, 10

Lipid to carbohydrate ratio (g/g) 2.47 6

1
Units stated in parentheses in column 1
2
Source: 1: Uys (1989); 2: Machiels and Henken (1985); 3: Machiels and Henken (1987); 4: Degani, Ben-Zvi and Levanon
(1989); 5: Fagbenro, Nwanna and Adebayo (1999); 6: Ali (2001); 7: Pantazis (1999); 8: Ali and Jauncey (2005b); 9: Ali
and Jauncey (2005c); 10: Ali and Jauncey (2005a); 11: Balogun and Ologhobo (1989); 12: Heinsbroek, Van Thoor and
Elizondo (1990); 13: Yilmaz et al. (2006).

Specifically formulated commercial catfish feeds are only available in certain

African countries (i.e. Cameroon, Kenya, Nigeria, South Africa and Uganda) (Hecht,
2007). Extruded floating or slow-sinking pellets are currently only available in Nigeria,
South Africa and Uganda. Large-scale, intensive catfish farmers throughout the African
subcontinent are currently (2010) still not entirely satisfied with feed quality. It was not
possible to obtain a list of ingredients that are used for the feeds except from Ugachick,
which uses soy bean, maize, sunflower and wheat products, mukene (silver cyprinid,
Rastrineobola argentea) fishmeal, unspecified fish oil, vegetable oil and bone meal from
Nile perch skeletons. Ugachick in Uganda produces three different feeds for Clarias,
the proximate composition of which is shown in Table 7. Because of the low protein
content of the pre-starter and starter feeds, farmers in Nigeria and Uganda import these
feeds from South Africa and elsewhere for the early rearing stages until the fish reach
~12 g (Ayinla, 2007; Rutaisire, 2007; Akinwole and Faturoti, 2007).
472 On-farm feeding and feed management in aquaculture

TABLE 7
Proximate composition of catfish feeds in Uganda and South Africa
Crude Crude Crude Ash Ca P
protein lipid fibre (% max) (%) (%)
(% min) (% min - max) (% max)

Ugachick
39.9 8.5–9.5 5.5 7
(high protein, pre-starter)

NutriScience (pre-starter) 45 8 4 3 7

Ugachick (starter) 35 8–10 7 7

NutriScience (starter) 40 8 4 3 7

Ugachick Catfish (grower) 30 7–8.5 8 7

NutriScience (grower) 35 7 4 3 7

Source: Information provided by feed manufacturers.

3.2 Feeding in intensive pond farming

Under both of the two intensive pond farming conditions described in sections 3.1
and 3.2 below, profitability depends largely on applying the correct feeding strategy.
This implies that the feed satisfies the nutrient requirements of the fish at various sizes,
feeding the right pellet size for easy consumption and low wastage, feeding the correct
amounts and feeding at the right time(s) each day. When fish are fed correctly, growth
rates are good and uniform across the population, FCR are low and pond water quality
is optimized (Isyagi et al., 2009b).

3.2.1 Under flow-through conditions

Under intensive pond-farming conditions with water exchange, the fingerlings
(~5 g) are stocked into grow-out ponds (0.1 ha) at a density of 100 000/ha and
fed according to feeding response and the feeding schedule shown in Table 5.
At the beginning of the production cycle and for about two months, the fish are
fed three times per day, after which the frequency is reduced to twice a day. The
fish population has to be thinned out at regular intervals to maintain a maximum
biomass of not more than 40 tonnes/ha, at a water exchange rate of total pond
volume once every four days. Under the prevailing thermal conditions, the fish can
be harvested at 800 g after 10 months (Uys, 1989).

3.2.2 Without water exchange

The production capacity of ponds under static water management conditions is lower
than under flow-through conditions. Under static conditions, using a complete sinking
diet, the critical standing crop is around 15–20 tonnes/ha. The critical standing crop is
defined as the biomass of fish in the pond when the growth rate begins to slow (Isyagi
et al., 2009b). While these yield levels are approximately 50 percent of those achieved
under flow-through conditions, they are still extremely high. Isyagi et al. (2009b)
suggest that fish <10 cm (~6 g) should not be used for on-growing. To obtain a lower
mortality rate and a higher level of profitability, they recommend larger fingerlings
of around 15 g. Based on a projected harvest weight of 800 g, 24 750 (15 g) fish are
required per hectare. This allows for a 10 percent mortality rate during the grow-
out period. Isyagi et al. (2009b) provide further detailed accounts of the feed choice,
management of feed conversion ratios (FCR), record keeping and economic analyses
of various feeding strategies under static pond-farming conditions in Uganda. Feed
costs make up approximately 60–70 percent of variable costs. Moreover, they (op cit.)
provide a suite of recommendations for on-farm feed management to ensure freshness
and good fish health.
Feed management practices for North African catfish in sub-Saharan Africa 473

3.3 Feeding in small-scale farming

The farming of catfish by small-scale commercial farmers is becoming more popular
in many sub-Saharan countries, including Mozambique, Zambia, Malawi, Kenya
and Tanzania. Many of these farmers either cannot afford or do not have access
to commercial formulated feeds and hence are forced to use farm-made feeds. For
economic reasons, over 90 percent of the feeds used by fish farmers in sub-Saharan
Africa are wet or dry feeds made on the farm or by small-scale feed producers (Hecht,
2007). Wet or moist ingredients that are commonly used for catfish feeds in Africa and
Asia include chicken entrails, minced poultry farm mortalities, abattoir waste, butchery
sweepings, fish market waste (mainly fish entrails), maggots, termites, earthworms,
trash fish, hotel or restaurant kitchen waste and live juvenile tilapia. In most instances,
these ingredients are mixed with milled oilseed cakes (soy, cotton, sunflower, palm
kernel) and relatively inexpensive ingredients such as maize, wheat or rice bran and
dried brewery waste.
Many farmers use wet ingredients such as chicken offal as a stand-alone feed
and claim to achieve FCR of around 1.3:1 (Ayinla, 2007). Recent trials in Uganda
have found that the use of chicken offal results in a high fat content of the meat and
unacceptably high abdominal fat deposition (Matsiko and Mwanj, 2008). However, it
should be noted that fish with low abdominal fat content are least preferred in many
other regions in Africa. There is some evidence to suggest that juvenile catfish fed on
mixed moist feeds (34 percent moisture) have poorer performance indices (i.e. weight
gain, specific growth rate, FCR, protein efficiency ratio) than juveniles fed on a dry diet
using the same ingredients (Fagbenro, 1994; Fagbenro and Jauncey,1994; Fagbenro,
Jauncey and Krueger, 1997).
Several farm-made dry feeds are produced, and these range from formulated dry
mixes fed to the fish in punctured bags, to moist feed cakes or balls and dry pellets.
Preparation and manufacturing technologies are simple – raw materials are milled and
cooked (or precooked, e.g. soybeans), mixed into a dough with water and extruded
using a meat mincer. The ‘spaghetti-like’ product is sun-dried or dried in locally
manufactured driers and cut or crumbled into appropriate sizes. Anti-oxidants are
expensive and not commonly used by small-scale feed manufacturers, such that storage
of farm-made feeds is problematic. This is compounded by the absence of proper
feed drying facilities. Farmers are advised to mix and prepare only quantities that can
be used within a few days. Depending on feeding practices, FCR of 1.1:1 have been
reported for some farm-made feeds in Uganda and Kenya (Hecht, 2007).
Because of the high cost of vitamin and mineral premixes, farmers in Africa often
use chicken layers mash, which contains minerals and vitamins, as one of the primary
components of catfish feeds. In fact, pig and poultry feeds are also often used as
alternate feeds for North African catfish throughout the African subcontinent (Liti,
MacWere and Veverica, 2002; Nyandat, 2007). Alternative sources of minerals used in
Nigeria include bone meal and crushed periwinkle shells (Ayinla, 2007). Gabriel et al.
(2007) and Hecht (2013) provide succinct overviews of alternative ingredients that have
been tested as ingredients for farm-made catfish feeds in sub-Saharan Africa.

3.4 Feeding in intensive tank culture conditions

The on-growing of African catfish under intensive tank culture conditions is somewhat
different to pond culture (Oellerman, 1995; Oellermann and Hecht, 1996; Hecht,
Oellermann and Verheust, 1996; Akinwole and Faturoti, 2007; Ayinla, 2007). Early
fingerling, juvenile and grow-out production schedules are now fairly standardized
and are briefly summarized as follows. Post-hatchery fry are stocked at a density of
~25 000/m3 and on-grown to 4–5 g in ~30 days. At that stage, they are graded into three
size classes. The smallest fish are discarded. Fish with an average weight of 4–5 g are
then restocked into the tanks at ~8 500/m3 and reared to 11–12 g in ~30 days. Up to
474 On-farm feeding and feed management in aquaculture

this stage, the volume of the rearing tanks is generally between 0.5 and 4 m3, while the
tanks used for on-growing in Nigeria and South Africa range in size from 2 to 20 m3
and are most commonly made of bricks and mortar. The 11–12 g fish are then stocked
at 150 to 400 fish/m3 and reared to market size (420–>850 g). At average temperatures
of between 26 and 28 oC, the 12 g fish are grown to 850 g (using 30 percent protein
feeds) in approximately 154 days at FCR ranging from 1.1 to 1.6:1 (Oellerman and
Hecht, 1996; Akinwole and Faturoti, 2007). Depending on management, production
levels range between 107 and 385 kg/m3/6 month cycle (Oellerman and Hecht, 1996;
Akinwole and Faturoti 2007).
Based on information provided by Akinwole and Faturoti (2007), it would
appear that high-density tank farming in Nigeria using recirculating systems is still
experiencing some teething problems, and these seem to be related to maintenance of
good water quality. Nevertheless, comparison of production parameters shows that
Nigerian farm results are good and approach the experimental results in the Netherlands
(cf. Akinwole and Faturoti, 2007; Eding and Kamstra, 2002 and Eding et al., 2006).
Because of the high densities at which the fish are reared, it is preferable to feed the
fish five to six times per day. Under high-density tank farming conditions, daily ration
tables are at best only guidelines; because of the high cost of feed, farmers are advised
to monitor food consumption and to adjust rations on a daily basis depending on the
feeding intensity of the fish. A sample of fish is normally weighed on a weekly basis (or
every 10 days) to check feed consumption against recommended levels.
Further strategies to reduce feed costs should be developed. For example, Ali (2001)
found that alternating periods of restricted feeding (maintenance requirement) for
three days and appetite feeding for the following four days is a plausible way in which
to reduce feed input cost for this species. However, in a later study, Ali and Jauncey
(2004) showed that C. gariepinus only shows partial compensatory growth under
periods of restricted and satiation feeding.
Feeding of African catfish under intensive tank culture conditions can be done by
hand or can be mechanized using demand feeders or compressed air feed distributors.
Demand feeders under these conditions are not recommended, as the pendulum is in
continuous motion, which may lead to feed losses. Despite the switch to intensive tank
culture, very little is known about the way to manage and optimize feed use under
these conditions, and there is a clear need to investigate this.

4. CONCLUSIONS
The farming of African catfish in sub-Saharan Africa is a commercial activity (defined
as a national production of >1 500 tonnes per annum) in only three countries viz.
Nigeria, Uganda and Kenya. In all other countries that report production statistics, the
activity is best described as marginal. The reasons for this are varied. In South Africa,
market resistance is the only impediment; while in many other countries access to
information, high cost and availability of capital, inadequate supplies and services and
poor infrastructure seem to be the overriding reasons. The technology for the farming
of the species at various levels of intensity is, however, very well established. It appears
that Uganda has recently become the centre of catfish farming development for food
and for the live bait sector.
There have been significant improvements in the aquafeed sector since 2005
(Hecht, 2007), both in terms of quality and supply. Most countries in the region
have adequate resources to manufacture appropriate feeds, although the availability
and cost of fishmeal and soybean meal or oilseed cake are major constraints in most
countries (Hecht, 2007). The general paucity of good-quality aquafeeds in the region
is generally a factor of scale. Uganda provides the best example of this supposition.
In 2005, commercial aquafeeds were not available in Uganda (Rutaisire, 2007) but
now, because of the rapid increase in catfish production, all the necessary feeds (fry,
Feed management practices for North African catfish in sub-Saharan Africa 475

fingerling and on-growing feeds) inclusive of floating extruded feeds are manufactured
locally. It is however unknown what proportion of the Ugandan catfish production
is reared on formulated and/or farm-made feeds. Although it is reported that farmers
in Nigeria and Uganda use high-protein imported feeds during the nursery phase, it
is anticipated that more appropriate feeds will also soon be manufactured in all major
catfish-producer countries.
Commercial feeding schedules during the nursery and on-growing phases have
also been fairly standardized. Live food (mainly Artemia) is generally used for the
first five to six days of exogenous feeding, after which the juveniles are weaned onto
dry feeds. During the early and late on-growing phases in ponds, the fish are fed by
hand to satiation at the same place and time; this reduces wastage. Under high-density
tank farming conditions, the fish are either fed by hand or through the use of demand
feeders. The use of the latter is considered wasteful, and further research is required to
optimize feeding practices in tanks under high-density conditions.
Subsistence farming of catfish under polyculture or monoculture conditions using
single ingredient feeds will remain marginal and make insignificant contributions to
food security and national production figures.

References
Akinwole, A.O. & Faturoti, E.O. 2007. Biological performance of African catfish
(Clarias gariepinus) cultured in recirculating system in Ibadan. Aquaculture
Engineering, 36: 18–23.
Ali, M.Z. 2001. Dietary protein and energy interactions in African catfish Clarias
gariepinus (Burchell, 1822). Ph.D, Thesis. Stirling, University of Stirling, 273 pp.
Ali, M.Z. & Jauncey, K. 2004. Evaluation of mixed feeding schedules with respect to
compensatory growth and body composition in African catfish Clarias gariepinus.
Aquaculture Nutrition, 10: 39–45.
Ali, M.Z. & Jauncey, K. 2005a. Effect of dietary lipid to carbohydrate ratios on body
composition, digestive enzyme activities and blood plasma components in African
catfish Clarias gariepinus (Burchell, 1822). Journal of Aquaculture in the Tropics,
20(1): 57–70.
Ali, M.Z. & Jauncey, K. 2005b. Effect of dietary protein to energy ratio on body
composition, digestive enzymes and blood plasma components in Clarias gariepinus
(Burchell, 1822). Indian Journal of Fisheries, 52(2): 141–150.
Ali, M.Z. & Jauncey, K. 2005c. Approaches to optimizing dietary protein to energy
ratio for African catfish Clarias gariepinus (Burchell, 1822). Aquaculture Nutrition,
11: 95–101.
Almazán, R.P., Schrama, J.W. & Verreth, J.A.J. 2004. Behavioural responses
under different feeding methods and light regimes of the African catfish (Clarias
gariepinus) juveniles. Aquaculture, 231: 347–359.
Amisah, S., Adjei-Boateng, D. & Afianu, D.D. 2008. Effects of bamboo substrate
and supplementary feed on growth and production of the African catfish, Clarias
gariepinus. Journal of Applied Sciences and Environmental Management, 12(2): 2–28.
Appelbaum, S. & van Damme, P. 1988. The feasibility of using exclusively artificial
dry feed for the rearing of Israeli Clarias gariepinus (Burchell, 1822) larvae and fry.
Journal of Applied Ichthyology, 4(3): 105–110.
ASEAN. 1978. Manual on pond culture of penaeid shrimp. A project of the ASEAN
with the assistance of the FAO/UNDP South China Sea Fisheries Development and
Coordinating Programme, Manila. ASEAN National Coordinating Agency of the
Philippines, Ministry of Foreign Affairs. 141 pp.
Awaiss, A. & Kestemont, P. 1998. Feeding sequences (rotifer and dry diet), survival,
growth and biochemical composition of African catfish, Clarias gariepinus Burchell
(Pisces: Clariidae), larvae. Aquaculture Research, 29: 731–741.
476 On-farm feeding and feed management in aquaculture

Ayinla, O.A. 2007. Analysis of feeds and fertilizers for sustainable aquaculture
development in Nigeria. In M.R. Hasan, T. Hecht, S.S. De Silva & A.G.J. Tacon, eds.
Study and analysis of feeds and fertilizers for sustainable aquaculture development,
pp. 453–470. FAO Fisheries Technical Paper No. 497, Rome, FAO. 510 pp.
Balogun A.M. & Ologhobo, A.D. 1989. Growth performance and nutrient utilization
of fingerling Clarias gariepinus (Burchell) fed raw and cooked soybean diets.
Aquaculture, 76: 119–126.
Bok, A.H. & Jongbloed, H. 1984. Growth and production of sharptooth catfish,
Clarias gariepinus (Pisces: Clariidae), in organically fertilized ponds in the Cape
Province, South Africa. Aquaculture, 36: 141–155.
Boyd, C.E. 1982. Water quality management for pond fish culture. Developments in
Aquaculture and Fisheries Science, 9. Amsterdam, Elsevier Scientific Publishing
Company. 318 pp.
De Graaf, G. & Janssen, H. 1996. Artificial reproduction and pond rearing of African
catfish, Clarias gariepinus in Sub-Saharan Africa. FAO Fisheries Technical Paper
No. 362. Rome, FAO. 73 pp.
De Kimpe, P. & Micha, J.C. 1974. First guidelines for the culture of Clarias lazera in
Central Africa. Aquaculture, 4: 227–248.
Degani, G., Ben-Zvi, Y. & Levanon, D. 1989. The effect of different protein levels
and temperatures on feed utilization, growth and body composition of Clarias
gariepinus (Burchell 1822). Aquaculture, 76: 293–301.
Eding, E.H. & Kamstra, A. 2002. Netherlands farms tune recirculation systems to
production of varied species. Global Aquaculture Advocate, 5: 52–54.
Eding, E.H., Kamstra, A., Verreth, J.A.J., Huisman, E.A. & Klapwijk, A. 2006.
Design and operation of nitrifying trickling filters in recirculating aquaculture: A
review. Aquaculture Engineering, 34: 234-260.
El Bolock, A.R. 1973. Breeding and culture of Clarias. FAO Aquaculture Bulletin, 5: 1–4.
Fagbenro, O.A. 1994. Studies on the use of fermented fish silage in diets for juvenile
tilapia, Oreochromis niloticus and catfish, Clarias gariepinus. Ph.D. Thesis. Stirling,
University of Stirling. 200 pp.
Fagbenro, O.A. & Jauncey, K. 1994. Growth and protein utilisation by juvenile catfish
(Clarias gariepinus) fed moist diets containing autolysed protein from stored lactic
acid fermented fish silage. Bioresource Technology, 48: 43–48.
Fagbenro, O., Jauncey, K. & Krueger, R. 1997. Nutritive value of dried lactic acid
fermented fish silage and soybean meal in dry diets for juvenile catfish, Clarias
gariepinus (Burchell, 1822). Journal of Applied Ichthyology, 13(1): 27–30.
Fagbenro, O.A., Nwanna, L.C. & Adebayo, O.T. 1999. Dietary arginine requirement
of the African catfish, Clarias gariepinus. Journal of Applied Aquaculture, 9(1):
59–64.
FAO. 2012. Fishstat Plus, Vers. 2.32. Rome, FAO. (available at www.fao.org/fishery/
statistics/software/fishstat/en).
Gabriel, U.U., Akinrotimi, O.A., Bekibele, D.O., Onunkwo, D.N. & Anyanwu, P.E.
2007. Locally produced fish feed: potentials for aquaculture development in Sub-
Saharan Africa. African Journal of Agricultural Research, 2(7): 287–295.
Haylor, G.S. 1993. Controlled hatchery production of African catfish, Clarias
gariepinus (Burchell): an overview. Aquaculture and Fisheries Management, 24:
245–252.
Hecht, T. 1996. An alternative life history approach to the nutrition and feeding of
Siluroidei larvae and early juveniles. Aquatic Living Resources, 9(5): 121–133.
Hecht, T. 2007. Review of feeds and fertilizers for sustainable aquaculture development
in Sub-Saharan Africa. In M.R. Hasan, T. Hecht, S.S. De Silva & A.G.J. Tacon, eds.
Study and analysis of feeds and fertilizers for sustainable aquaculture development,
pp. 77–110. FAO Fisheries Technical Paper No. 497. Rome, FAO. 510 pp.
Feed management practices for North African catfish in sub-Saharan Africa 477

Hecht, T. 2013. Species profile: North African catfish, Clarias gariepinus (Burchell,
1822). In Aquaculture Feed and Fertilizer Resources Information System (www.fao.
org/fishery/affris/en/org/fishery/affris/en/). Rome, FAO.
Hecht, T. & Appelbaum, S. 1988. Observations on intraspecific aggression and coeval
sibling cannibalism by larval and juvenile Clarias gariepinus (Clariidae: Pisces)
under controlled conditions. Journal of Zoology, 214: 21–44.
Hecht, T. & Pienaar, A.G. 1993. A review of cannibalism and its implications in fish
larviculture. Journal of the World Aquaculture Society, 24: 246–261.
Hecht, T., Oellermann, L. & Verheust, L. 1996. Perspectives on clariid catfish culture
in Africa. Aquatic Living Resources, 9(5): 197–206.
Hecht, T., Uys, W. & Britz, P.J., eds. 1988. The culture of sharptooth catfish, Clarias
gariepinus in southern Africa. South African National Scientific Programmes
Report No. 153. Pretoria, Council for Scientific and Industrial Research. 133 pp.
Heinsbroek, L.T.N., Van Thoor, R.M.J. &. Elizondo, L.J. 1990. The effect of feeding
level on the apparent digestibilities of nutrients and energy of a reference diet for
the European eel, Anguilla anguilla and the African catfish, Clarias gariepinus. In
M. Takeda & T. Watanabe, eds. The current status of fish nutrition in aquaculture,
pp. 175–188. Proceedings of the Third International Symposium on Feeding and
Nutrition in Fish, August 28 – September 1, 1989, Toba, Japan. Tokyo, Tokyo
University of Fisheries. 469 pp.
Hepher, B. 1963. Ten years of research in fish pond fertilization in Israel. 11. Fertilizer
dose and frequency of fertilization. Bamidgeh, 15(4):78–92.
Hepher, B. 1967. Some limiting factors affecting the dose of fertilizers added to
fishponds, with special reference to the Near East. In T.V.R. Pillay, ed. Proceedings
of the FAO World Symposium on Warm-water Pond Fish Culture, Rome, Italy,
18–25 May 1966, FAO Fisheries Report No. 44, Vol. 3: 1–6.
Hepher, B. & Pruginin, Y. 1981. Commercial fish farming. New York, John Wiley &
Sons, (Publishers) Inc., 261 pp.
Hey, D. 1941. Practical freshwater fish culture. Cape Town, Cape Nature Conservation,
118 pp.
Hogendoorn, H. 1979. Controlled propagation of the African catfish, Clarias lazera
(C. and V.), 1: reproductive biology and field experiments. Aquaculture, 17: 323–333.
Hogendoorn, H. 1980. Controlled propagation of the African catfish, Clarias lazera
(C. and V.). III: feeding and growth of fry. Aquaculture, 21: 233–241.
Hogendoorn, H. 1981. Controlled propagation of the African catfish, Clarias lazera
(C. & V.), IV: effect of feeding regime in fingerling culture. Aquaculture, 24: 123–131.
Hogendoorn, H., Jansen, J.A.J., Koops, W.J., Machiels, M.A.M., van Ewijk, P.H. &
van Hees, J.P. 1983. Growth and production of the African catfish, Clarias lazera
(C. & V.). II. Effects of body weight, temperature and feeding level in intensive tank
culture. Aquaculture, 34: 265–285.
Isyagi, N.A., Veverica, K.L., Asiimwe, R. & Daniels, W.H. 2009a. Manual for the
commercial pond production of the African catfish in Uganda. Auburn University,
Alabama. 238 pp. (available at: www.ag.auburn.edu/fish/international/uganda/).
Isyagi, N., Atukunda, G., Aliguma, L., Ssebisubi, M., John, W., Kubiriza, G. &
Mbulameri, E. 2009b. Assessment of national aquaculture policies and programmes
in Uganda. SARNISSA EC FP7 Project. Contract Number 213143, Stirling, United
Kingdom, University of Stirling. 79 pp. (available at: www.sarnissa.org).
Janssen, J.A.L. 1985. Elevage du poisson-chat africain Clarias lazera (C&V) en
République Centraficaine. III. Alevinage et grossissement en étangs. FAO projet
GCD/CAF/007/NET. Document Technique No. 22, 41 pp. Rome, FAO.
Kerdchuen, K. 1992. L’alimentation artificielle d’un silure africain, Heterobranchus
longifilis (Teleostei: Clariidae). Incidence du mode d’alimentation et première
estimation des besoins nutritionnels. These de Doctorat de l’Universite Paris 6, 182 pp.
478 On-farm feeding and feed management in aquaculture

Liti, D., MacWere, O.E. & Veverica, K.L. 2002. Growth performance and economic
benefits of Oreochromis niloticus/Clarias gariepinus polyculture fed on three
supplementary feeds in fertilized tropical ponds. (Available at: http://pdacrsp.
oregonstate.edu/pubs/technical/19tchtml/9FFR2.html).
Machiels, M.A.M. & Henken, A.M. 1985. Growth rate, feed utilization and energy
metabolism of the African catfish, Clarias gariepinus (Burchell, 1822), as affected
by dietary protein and energy content. Aquaculture, 44: 271–284.
Machiels, M.A.M. & Henken, A.M. 1987. A dynamic simulation model for growth of
the African catfish, Clarias gariepinus (Burchell, 1822) II. Effect of feed composition
on growth and energy metabolism. Aquaculture, 60: 33–53.
Matsiko, S.D. & Mwanj, M.T. 2008. The current status of catfish culture and research
in Uganda. In R.W. Ponzoni & N.H. Nguyen, eds. Proceedings of a Workshop on
the Development of a Genetic Improvement Program for African Catfish Clarias
gariepinus, pp. 57–63. Penang, WorldFish Center Conference Proceedings No. 1889.
Micha, J.C. 1971. Pond culture of African catfishes. FAO Aquaculture Bulletin, 3: 6.
Micha, J.C. 1972. Induced breeding of Clarias spp. FAO Aquaculture Bulletin, 4: 3–4.
Nwachukwa, V.N. 1999. Periphyton fauna as an alternative live food in the rearing of Clarias
gariepinus (Burchell) fry. Israeli Journal of Aquaculture - Bamidgeh, 51(4): 169–171.
Nyandat, B. 2007. Analysis of feeds and fertilizers for sustainable aquaculture
development in Kenya. In M.R. Hasan, T. Hecht, S.S. De Silva & A.G.J. Tacon, eds.
Study and analysis of feeds and fertilizers for sustainable aquaculture development,
pp. 423–436. FAO Fisheries Technical Paper No. 497. Rome, FAO. 510 pp.
Oellermann, L.K. 1995. A comparison of the aquaculture potential of Clarias gariepinus
(Burchell, 1922) and its hybrid with Heterobranchus longifilis Valenciennes, 1840 in
Southern Africa. Ph.D. Thesis, Grahamstown, Rhodes University, 147 pp.
Oellermann, L.K. & Hecht, T. 1996. High density culture of hybrid catfish. In P. Cook
& W. Uys, eds. Proceedings of the Third Congress of the Aquaculture Association
of Southern Africa, Berg-en-Dal, Kruger National Park, 21–23 September 1994, pp.
116–128. Pretoria, Aquaculture Association of Southern Africa. 154 pp.
Pantazis, P.A. 1999. Nutritional studies in the African catfish, Clarias gariepinus
(Burchell, 1822). Ph.D. Thesis. Stirling, University of Stirling, 192 pp.
Richter, C.J.J. 1976. The African catfish, Clarias lazera (C. and V.), a new possibility
for fish culture in tropical regions? In E.A. Huisman, ed. Aspects of fish culture
and fish breeding. Miscellaneous Papers of the Landbouwhogeschool, Wageningen
(Netherlands), 13: 51–71.
Rutaisire, J. 2007. Analysis of feeds and fertilizers for sustainable aquaculture
development in Uganda. In M.R. Hasan, T. Hecht, S.S. De Silva & A.G.J. Tacon, eds.
Study and analysis of feeds and fertilizers for sustainable aquaculture development,
pp. 471–487. FAO Fisheries Technical Paper No. 497. Rome, FAO. 510 pp.
Segner, H., Rosch, J., Verreth, J. & Witt. U. 1993. Larval nutritional physiology:
studies with Clarias gariepinus, Coregonus lavaretus and Scopthalmus maximus.
Journal of the World Aquaculture Society, 24: 121–133.
Stroband, H.W.J. & Kroon, A.G. 1981. The development of the stomach in Clarias
lazera and the intestinal absorption of protein macromolecules. Cell and Tissue
Research, 215(2): 397–415.
Tacon, A.G.J. 1987. The nutrition and feeding of farmed fish and shrimp – a training
manual. 2. Nutrient sources and composition. FAO Field Document No. 5, Brasilia,
Brazil. 129 pp.
Tacon, A.G.J. 1988. The nutrition and feeding of farmed fish and shrimp – a training
manual. 3. Feeding methods. FAO Field Document No. 7, Brasilia, Brazil. 208 pp.
Uys, W. 1984. Investigations into the dietary requirements of Clarias gariepinus larvae
(Pisces: Clariidae) and the formulation and manufacture of an artificial dry feed for use
in intensive larval rearing. M.Sc. Thesis. Grahamstown, Rhodes University. 122 pp.
Feed management practices for North African catfish in sub-Saharan Africa 479

Uys, W. 1989. Aspects of the nutritional physiology and dietary requirements of juvenile
and adult sharptooth catfish, Clarias gariepinus (Pisces; Clariidae). Ph.D. Thesis.
Grahamstown, Rhodes University. 190 pp.
Uys, W. & Hecht, T. 1985. Evaluation and preparation of a suitable dry feed and
optimal feeding frequency for the primary nursing of Clarias gariepinus larvae
(Pisces: Clariidae). Aquaculture, 47: 173–183.
Verreth, J. & Den Bieman, H. 1987. Quantitative feed requirements of African catfish
(Clarias gariepinus Burchell) larvae fed with decapsulated cysts of Artemia. 1. The
effect of temperature and feeding level. Aquaculture, 63: 251–267.
Verreth, J. & Van Tongeren, M. 1989. Weaning time in Clarias gariepinus (Burchell)
larvae. Aquaculture, 83: 81–88.
Verreth, J., Storch, V. & Segner, H. 1987. A comparative study on the nutritional
quality of decapsulated Artemia cysts, micro-encapsulated egg diets and enriched
dry feeds for Clarias gariepinus (Burchell) larvae. Aquaculture, 63: 269–282.
Verreth, J., Eding, E.H., Rao, G.R.M., Huskens, F. & Segner, H. 1993. A review
of feeding practices, growth and nutritional physiology in larvae of the catfishes
Clarias gariepinus and Clarias batrachus. Journal of the World Aquaculture Society,
24: 135–144.
Verreth, J.A.J., Torreele, E., Spazier, E., van der Sluiszen, A., Rombout, J.H.W.M.,
Booms, R. & Segner, H. 1992. The development of a functional digestive system in
the African catfish Clarias gariepinus (Burchell). Journal of the World Aquaculture
Society, 23: 286–298.
Viveen, W.J.A.R., Richter, C.J.J., Van Oordt, P.G.W.J., Janssen, J.A.L. & Huisman,
E.A. 1985. Practical manual for the culture of the African catfish (Clarias gariepinus).
The Hague, The Netherlands Ministry for Development Cooperation, Section for
Research and Technology. 128 pp.
Yilmaz, E., Akyuart, I., Mutlu, E. & Akyurt, I. 2006. Effects of energetic diets on
growth, blood chemistry, and liver pathology of African catfish, Clarias gariepinus
(Burchell 1822). Israeli Journal of Aquaculture – Bamidgeh, 58(3): 191–197.