Journal of Zoology - December 1978 - Willoughby - The Ecology of The Catfish Clarias Gariepinus and Clarias Ngamensis in

J . Zool., Lond.
(1978) 186, 507-534
The ecology of the catfish Clarias gariepinus and Clarias ngamensis

in the Shire Valley, Malawi
N . G. W I L L O U G H BAYN D D. T W E D D L E
Fisheries Department, P.O. Box 593, Lilongwe, Malawi
(Accepted 11 April 1978)
(With 2 plates and 15 figures in the text)
The ecology of two commercially important catfish species, CIarias gariepinus and C .
ngamensis, was examined in the Elephant Marsh, southern Malawi. Abundance, distribution,
and movements within the marsh system were studied. Growth rates were determined from
rings on the vertebrae, with corroborative evidence from length frequency analyses and
tagging studies. Breeding seasons, length and age at maturity, and fecundity were deter-
mined for both species. Stomach contents were examined and the food preferences of the
two species were compared.
Contents
Page
Introduction.. . . . . . . . . . . .. .. .. .. .. 508
Description of study area . . . . . . . . .. .. .. .. .. 509
Methods .. .. .. .. . . . . .. .. .. .. .. 509
Capture methods .. .. .. .. .. .. .. .. .. 509
Movement .. .. .. . . . . .. .. .. .. .. 510
Ageand growth . . . . . . .. .. .. .. .. .. 510
Length-weight relationship . . . . . . .. .. .. .. .. 510
Breeding . . . . . . . . . . .. .. .. .. .. 510
Feeding.. .. .. .. . . . . .. .. .. .. .. 511
Results .. .. .. .. . . . . .. .. .. .. .. 51 1
Abundance and distribution . . . . .. .. .. .. .. .. 51 1
Migration-C. gariepinus . . .. .. .. .. .. .. .. 513
Age and growth . . . . .. .. .. .. .. .. .. 513
Length frequency analysis .. .. .. .. .. .. .. 513
C . gariepinus .. . . . . .. .. .. .. .. 513
C . ngamensis . . . . . . .. . . . . .. .. 516
Growth rates calculated from vertebrae .. .. .. .. .. 516
C . gariepinus .. . . . . .. .. .. .. .. 516
C. ngamensis .. . . . . .. .. .. .. .. 517
Growth of tagged fish-C. gariepinus . . .. .. .. .. .. 518
Length-weight relationship . . . . .. .. .. .. .. .. 519
Breeding . . . . . . . . . . .. .. .. .. .. 521
Sex ratios. . . . .. . . . . .. .. .. .. .. 521
Length at maturity .. .. .. .. .. .. .. .. 521
Fecundity. . .. .. .. .. .. .. .. .. .. 522
Breeding seasons and sites . . . . .. , . .. .. .. 522
Food and feeding .. . . . . .. .. .. .. .. .. 526
C. gariepinus . . . . . . . . .. . . . . .. .. 526
C. ngamensis . . . . . . . . .. ). .. .. .. 526
Discussion . . . . . . . . . . . . .. ,. .. .. .. 521
Summary . . . . . . . . . . . . .. . . . . .. .. 532
References . . . . . . . . . . . . .. . . . . .. .. 533
507
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508 N . G . WILLOUGHBY A N D D. T W E D D L E
Introduction
The catfish Clarias gariepinus (Burchell) (= C. mossambicus Peters)* and C. ngarnensis
Castelnau are of considerable importance in the traditional fishery of the River Shire in
southern Malawi, where C. gariepinus makes up over 50% and C. ngamensis 20% of the
annual catch of 12-14,000 tons. C. gariepinus is commercially important in much of east,
central and southern Africa.
Information on the biology of C. gariepinus has been given by several workers (Green-
wood, 1955; Corbet, 1961; Groenewald, 1964; Kirk, 1967; Hastings, 1973 (unpublished
report); van der Waal, 1974; van der Waal & Schoonbee, 1975), but despite its importance
no full account of its ecology in a single area has been published. Apart from brief notes
written by Jackson (1961), Jubb (1967) and Bell-Cross (1974) no published work on C.
ngamensis has been located. This paper presents information on all aspects of the ecology
of both species in the Elephant Marsh area of the River Shire.
0,
I
/
Chilwa
1
I
I
I
I
I
,' I
I
I
I
I
.- '.
\
I
FIG. 1 . The Shire Valley and its associated swamps, southern Malawi.
* In this paper we follow Jubb (1967) in considering Clurius guriepinus to be the senior synonym of Clurius
mossumhicus.
ECOLOGY O F THE CATFISH 509
Description of study area

The marshes in the lower valley of the R. Shire cover an area of approximately 650 km2
(Fig. 1). There are two main areas, the Elephant Marsh (500 km2) to the north, and the
Dinde Marsh (1 50 kmz) to the south. These are joined by a predominantly riverine section,
although small areas of marsh occur on the eastern side of the R. Shire in Mocambique.
The marshes are subject to considerable changes in water levels as a result of two factors;
the local rains, and a barrage at Liwonde which was built to regulate the level of Lake
Malawi and consequently also regulates the flow of the R. Shire. The marshes are relatively
dry between September and November, and fill during December (Fig. 2).
1974 ' I975

I
1976
FIG.2. Seasonal variation in the water level (height in m above Shire Valley datum) and rainfall at Chiromo
August 1974July 1976.
Three areas in the Elephant Marsh were regularly sampled. At Chiromo, the confluence
of the Ruo and Shire rivers, the habitat is mainly riverine with the vegetation along the
banks dominated by grasses of the genus Echinochloa. At Mchacha there are large open
lagoons usually less than 4 m deep containing dense beds of Ceratophyllum and Nymphaea.
The lagoons are separated by areas dominated by Phragmites, but are interconnected by
small channels and the whole area is normally subject to low rates of flow. At Ndombo,
on the eastern shore of the Elephant Marsh both riverine and lagoon habitats are found,
the dominant emergent vegetation being Typha, with C'eratophyllum abundant in the
lagoons.
Methods
Capture methods
Specimens were caught between August 1974 and July 1976 using graded fleets of multifilament
nylon gill nets. The original fleets had stretched mesh sizes of 5.1, 6.4, 7.6, 8.9, 10.2, 12.7, 15.3,
510 N . G. W I L L O U G H B Y A N D D. T W E D D L E
and 17.9 cm (2, 24, 3, 34, 4, 5, 6 and 7 in). The 12.7, 15.3 and 17.9 cm nets were not set after
July, 1975, as in the entire year in which they were set they caught only one fish. In April 1975,
2.5 and 3-8 cm nets (1 and 19 in) were added to the fleets, but these also caught few Clarias.
The 5.1-17.9 cm nets were 45.8 m (50 yds) long and 1.2 m (4 ft) deep when mounted by the half.
The 2.5 and 3.8 cm nets were the same depth but only half the length.
The nets were set at each of three stations in the Elephant Marsh for 12 nights each month.
They were set during the afternoon and usually lifted the following morning, although on certain
occasions they were left throughout the day if set unusually far from the station. The extended
period of these sets had little effect on the catch rate of Clarias as the species are largely nocturnal.
At Chiromo nets were set in the stream parallel to the current and close to dense banks of
vegetation; at Mchacha they were set in more static water, while those at Ndombo sampled both
riverine and lagoonal habitats. In addition to the fish caught in gill nets, specimens caught by
other methods were examined to supplement the data on certain aspects of the biology of the
fish. All Clarias were measured to the nearest mm total length and weighed to the nearest gram
for fish below 200 g and the nearest 25 g for larger specimens.
Movement
Tagging studies were carried out t o investigate movement of C. gariepinus within the marsh.
Specimens were caught at several sites using electrofishing apparatus and numbered plastic flag
tags were attached to their lower jaws using galvanized wire. They were released at the capture
sites.
Age and growth

Preliminary examination of skeletons of both species revealed consistent zones on the vertebrae.
Vertebrae were therefore collected from specimens caught at each station every month between
August 1974 and July 1975. They were prepared by boiling the head section, then removing the
first four vertebrae from immediately behind the vertebral complex at the base of the skull.
Cooked flesh was removed by washing and no further treatment was necessary. The rings
consisted of alternate hyaline and opaque zones which were quite distinct using either reflected
or transmitted light. Measurements were made along the radius from the centre to the dorsal
edge of the centrum. The distance from the centre to the start of each hyaline zone was measured
using an eyepiece graticule in a stereo dissecting microscope which had been calibrated against
a mm scale. The third and fourth vertebrae were used as they were more regular in shape than
the first two.
Length-weight relationship
Total lengths and weights of 1888 specimens of C. gariepinus and 879 of C. ngamensis (50 of
each cm size group if available) were used to determine the length/weight relationships. Sexes
were not separated as no significant differences were observed between them, and data were
taken over a whole year to eliminate differences due to changes in condition.
Breeding
The sexes of all fish and their states of maturity were noted. Maturity for females was defined
as when the oocytes had fully developed, and for males when the testes exuded milt. Ripe ovaries
were removed, weighed and stored in Gilson’s Fluid. The eggs were later separated from the tissue
and dried at room temperature. The fecundity of each fish was determined by weighing the total
number of eggs in the ovaries, followed b y two sub-samples, each of 250 eggs, using the method
outlined by Simpson (1959).
ECOLOGY O F T H E CATFISH 511
Feeding
Stomachs were collected from both species caught in the nets between August 1974 and July
1975. Some stomachs of C. gariepinus caught on flooded farmland in May 1976, and of others
caught in traps in the marsh were also examined. All the stomachs were preserved in 4 % formalin
and their contents later removed, weighed and examined under a dissecting microscope. The
contents were separated into several categories, three of which, plant detritus, humus and mud
were determined by particle size. Plant detritus was considered to be dead plant material which
would not pass through an Endecott sieve of 425 pm pore size; humus was that material retained
between sieves of 420pm afid 125pm pore size, and mud was the residue. The item which
contributed the greatest mass to the stomach contents was termed the dominant item for that
stomach.
I
TABLE
Numbers and weights of Clarias caught at sampling stations (August 1974-July 1976)
Sampling O/
,,Total
, no. of Weight 7; Total weight of
station Number fish at station (kg) fish at station
C. gariepinus
Chiromo 998 12$4 429.6 41 %
Mch acha 2275 22 % 886.3 53 %
Ndombo 2802 29 % 942.1 60 %
Total 6075 - 2258.0 -
C. ngamensis
Chiromo 412 5% 146.0 15%
Mchacha 1393 14% 413.7 26 %
Ndombo 1183 13% 289.2 19%
Total 2988 - 848.9 -
Results
Abundance and distribution
The total numbers and weight of Clarias caught during the 24 month sampling period
are shown in Table I. The importance of both species to the traditional fishery are obvious
from these figures. Although fishermen do not use the full range of mesh sizes used in the
research programme, the proportion of C. gariepinus in the 6-4 cm to 8.9 cm mesh nets
commonly used is similar and C . gariepinus constitutes slightly more than 50% of the
traditional fishery catch for all gears. C. gariepinus were most abundant at Ndombo and
C. ngamensis at Mchacha. Both species were least abundant at Chiromo.
Seasonal variations in the numbers and weights are shown in Fig. 3. In both species
peak catches occurred in September 1974 and August 1975 and also from December 1974
to March 1975 and in January 1976. C. ngamensis catches in the 1975176 wet season
were considerably less than those in the previous year.
There were significant reductions in catches of both species in November each year and
also in May and June, with the exception of C. ngamensis in 1976, where the catch recovered
in these months from a very low level in April.
512 N . G. W I L L O U G H B Y A N D D. T W E D D L E
-
-m
s
500 -
400-
'0 '. .'.,,._.-.
Individual wght (Snaothed m n s )
\
./.-.-.
-/ .'-.-...-._.~.
1
A S O N D J F M A M J J A S O N D J F M A M J J
1974 1975 1976
b t a l number and weight

0-0 Number
lndividuol weigMs (Smoothed means)
A S O N D I J F M A M J J A S O N D J F M A M J J
1974 1975 I976

Fig. 3. Seasonal variation in total number, total weight and mean individual weights of (a) C . guriepinus and
(b) C . ngamensis caught at sampling sites.
The mean weights of the individual fish caught each month are also shown in Fig. 3.
The mean weights of the specimens caught at the stations over the whole sampling period
were :
C. gariepinus: Chiromo 430 g, Mchacha 390 g, Ndombo 340 g
C. ngamensis: Chiromo 360 g, Mchacha 270 g, Ndombo 230 g
Migra tion-C. gaviepinus

There were 40 returns from 238 tagged fish within 300 days of the original tagging.
These returns showed that 10 fish had moved upriver, 5 downriver, and 25 were recovered
very close to the tagging site. The only fish which had moved rapidly away from the
tagging site was one which moved down-river at more than 1 km/day for 42 days. These
movements are shown graphically in Fig. 4. There was no correlation between the size of
a fish and the distance which it moved.
I 120
t . . .
I I I
100 200 300
Time belween logging and recapture (days)
FIG.4. Movement of tagged C. gariepinus, indicating the distance each tagged fish moved upstream or down in
the period between release and recapture.
Age and growth

Length frequency analysis
C. gariepinus. Data for the year April 1975 (when 2.5 and 3.8 cm nets were introduced)
to March 1976 are shown (Fig. 5) in 2 cm length groups, with, e.g. the 21 cm group including
fish from 20.0-21.9 cm. The sexes were combined as there was no noticeable difference in
length distribution or growth in the first three years. It is evident that there is either
considerable variation in the growth rates of individual fish, or a prolonged breeding
season, and that any conclusions on growth rates must be considered tentative and
dependent on confirmation from other sources.
Figure 6 gives the net mesh selectivity for this species, and shows a similar skewed
pattern to that found by Gulland & Harding (1961), with closely corresponding peaks for
7.6 cm and 10.2 cm mesh nets. Correction for net selectivity was not considered necessary
because of the small mesh increments (1.3 cm) and corresponding closeness of the modes
for each net.
514 N . G . WILLOUGHBY A N D D . T W E D D L E
2 3
April '75
N.227
10- May
0 - - - I I5
10-
0-
10-
25 35
-
June
July
66
0- - 128
10- Dee
288
0
21 31
Jan '76
10-
0. -
I I 409
10- Feb
0-
_ I -
- 262
March
Length (cm 1
Frc. 5. Length frequency distribution of C . gariepinus, both sexes combined. N=numbers of fish in monthly
sample. The figures above the histograms indicate the suggested age in years of each peak in the histograms.
The peak of the breeding season occurs in December (see next section), and therefore
the first mode in the length frequency data for April 1975, at 19 cm, may be tentatively
assumed to be composed of fish of approximately 1.3 years old. The next peak, at approxi-
mately 29 cm, would then represent 2.3 year old fish. The assumed age for each peak is
given for subsequent months on Fig. 5. The indications are that 2-0 year old fish were 30
cm in length and 3.0 year olds about 38 cm in December. Fish of approximately one year
old began to appear in catches during January and February.
The length distribution of small C. gariepinus caught while electrofishing in water less
than 25 cm deep is shown in Fig. 7. Assuming that spawning occurs at the end of
December, they reach 11 cm in four months and 15-16 cm by the cold season at six months
old. Four larger fish of 26-31 cm were also taken in the shallow water, and their lengths
correspond with the modal length of 26 cm for 1.5 year old fish in July 1975 (Fig. 5). The
FIG.6. Length composition of catches of C. guriepinus by gill nets of various mesh sizes. Stretched mesh size
515
35
7.7.76(kngochi)
13.4.76(Mchocha)
30
15.6.76(Chiromo)
3 8crn
6 4crn
2 5 cm
I
7 6 crn
5 Icrn
0 9 cm
10 2 cm
I
70
I
60
I
25
I
I
ECOLOGY O F T H E CATFISH
50
Length d fish (crn)

I
20
I
I
40
I
3
-
A
5
1
50
/'-
10
I
I
m
given at right of figure.

1
-
-
0.
5-
5.
Oh
I-
0
s
E
n
b
c
Lc
0
.4-
516 N . G . W I L L O U G H B Y A N D D. T W E D D L E
Mangochi swamps from which some of the fish were taken are not in the lower part of
the Shire Valley, being 400 m higher up the valley, but they form a similar environment
with a similar flooding regime, so it is believed that the growth of the C. gariepinus, at
least in the juvenile stages, is comparable in the two areas.
C. ngamensis. It was not possible to make any growth estimates from the available
length frequency data for this species.
Growth rates calculated from vertebrae

C. gariepinus. Opaque and hyaline zones had to be consistent on all four vertebrae to
be considered true rings. Occasionally false rings were present, distinguishable by their
incomplete or indistinct appearance. The vertebrae from 150 out of the total of 628 fish
examined were discarded as unreadable, and those from the remaining 255 males and 223
females were used for growth determinations.
Hyaline zones were found at the edges of the centra of some vertebrae in June and
July, and by September the hyaline rings could be seen near the edges of the majority of
vertebrae, so it appears that the rings are laid down during the cold season, May-August.
The centres of the vertebrae were variable in appearance. Most were hyaline, and in
these the outer edge of this zone was measured as the first ring and considered to be the
size of the centrum at the end of the first cold season of life, i.e. at an age of approximately
PLATEI. Vertebrae from a male C. guriepinus 45 cm in length and estimated to be 3.8 years old.
ECOLOGY OF T H E CATFISH 517
0.7 years. For those vertebrae with opaque centres the distance to the first hyaline zone
was measured, this being the size at approximately 0.5 years. These assumptions were
corroborated by examination of vertebrae of small fishes caught under natural conditions
and considered, on available evidence, to be in their first year. Plate I shows vertebrae
from a male fish 45 cm long, caught in October, and estimated to be 3.8 years old. The
ring formed in the cold season at age 3.5 can be seen at the edge of the centrum.
The relationship between the total length in cm (L) and the vertebral radius in cm (R)
was best described by the equation log L = 0.6876 log R + 1.9412 with a correlation co-
efficient of 0.89. The lengths when the rings were laid down in each fish were then calcu-
lated using the method described by Le Cren (1947). The average lengths for each ring in
Age (years)
FIG.8. Growth curves of (a) C. guriepinus and (b) C. ngumensis calculated from vertebrae. Numbers with the
points indicate the number of vertebrae involved in the calculations. Top curve, males; bottom curve, females.
both sexes are shown in Fig. 8. The figure for the first ring is an average for length in the
cold season, but for other rings the sizes shown are those at the beginning of the cold
season, when the hyaline zones start to be laid down. Using the data in Fig. 8, von
Bertalanffy growth curves were calculated for each sex following the method outlined by
Ricker (1975). The maximum length ( Lm) for males was calculated to be 139.4 cm and for
females 78.9 cm with growth constants K (a measure of the rate at which L approaches
L cc) = 0.0854 and 0.1668 respectively.
Both sexes reach approximately 20 cm in their first year, 30 cm in the second and 38 cm
in the third. Females grow more slowly thereafter, and the majority of large specimens are
male. The largest specimen recorded was a male of 100 cm in length and 8.5 kg in weight.
C. ngamensis. Vertebrae from 392 fish were examined of which 83 were discarded as
unreadable. Those from the remaining 167 males and 142 females were used for growth
determinations.
The rings had the same appearance as those of C. gariepinus, and the same procedure
was followed.
In July a number of vertebrae were seen to have hyaline zones right at the edge of the
centrum, and by September-October this ring was distinguishable on the majority of the
518 N. G . WILLOUGHBY A N D D. TWEDDLE
vertebrae (Fig. 9). It therefore appears that this ring is laid down in the cold season,
May-August. Figure 9 also suggests that rapid growth occurs in the warmest months,
from December to April.
Plate I1 shows a vertebra from a female fish 45.7 cm long, caught in June 1976, and
estimated to be 3-5 years old. No peripheral hyaline zone is yet distinguishable.
L Aug '74
f Ln,
Feb
Mar
I--%--
June
Jan '75 July
Fig. 9. Distance of last hyaline zone from edge of centrum of vertebra, on a monthly basis, for C. ngurnensis.
The relationship between total length (L) and vertebral radius (R)was log L=0.7045
log R + 1-9416with a correlation coefficient of 0-93. The average lengths for each ring are
shown in Fig. 8. Von Bertalanffy growth curves were calculated and the maximum length
( L m) for males was calculated to be 73.1 cm and for females 69.8 cm, with growth constants
K = 0.1788 and 0.1779 respectively.
The largest specimen caught during the survey was a 55 cm male weighing 1.5 kg.
Growth of tagged fish-C. gariepinus

Four tagged fish were free for more than five months including the period of fastest
growth. Their growth increments are shown in Table 11. Three had shown good growth,
but the fourth had apparently grown only 1-5cm in length despite having put on 165 g
in weight. Errors in measurement are most likely to occur when handling the active fish
during tagging, and it is thought that an error may have occurred in this case, leading to
ECOLOGY OF THE CATFISH 519
PLATE11. Vertebra from a female C . ngamensis 45.7 cm in length and estimated to be 3.5 years old.
an underestimate of the growth in length of this specimen. The increase in length of the
other three fish is of the order one would expect from other evidence, and suggests that
the presence of the tag did not interfere unduly with normal feeding behaviour.
Length-weight relationship
The relationships between the total length in cm (L) and the weight in g ( W ) are
described by the equations :
C . gariepinus: log W = 3.136 log L-2.3396

C. ngamensis: log W= 3.2849 log L - 2.5429
T A B L EI1
Growth of tagged C. gariepinus, indicating increase in length in cm
during the period in which the Jish were at large hefween release
after tugging andrecapture
Length when Length when

Days free released (cm) recaptured (cm) Increment
292 28.0 34.0 6.0

273 39.0 40.5 I .5
233 35.0 40.0 5.0
172 51.0 57.0 6.0
520 N . G . WILLOUGHBY A N D D . TWEDDLE
T A B L EI11
Number of specimens of each sex of both species caught at each
station, to indicate the sex ratios
Sampling
station Males Females Ratio (Male/Female)
C . gariepinus
Chiromo 406 438 0.93 : 1 . 0 0
Mchacha 1144 971 1.18 : 1.00
Ndombo 956 899 1.06 : 1.00
Total 2506 2308 1.09 : 1.00
C. ngamensis
Chiromo] 205 157 1.31 : 1.00
Mchacha 81 1 691 1.17 : 1 . 0 0
Ndombo 46 1 380 1.21 : 1.00
Total 1477 1228 1.20 : 1.00
25
X : x X
X I
Length ( c m )
FIG.10. Average length at first maturity for C . gariepinus. The dashed line indicates the maximum percentage of
mature fish in any length group. The dotted line indicates 50% of this figure which is taken as the average length
at maturity.
Breeding
Sex ratios
The overall sex ratio and also the sex ratios of the fish at each of the three stations are
given in Table 111. There were slight differences between the sex ratios of fish Ianded at
the three sites, and the overall ratio is significantly different from unity, with more males
than females being caught for both species.
Length at maturity
Beverton & Holt (1957) defined the average weight at first maturity as being the weight
at which 50% of the individuals first reach maturity. Similarly average length at first
maturity may be defined as the length at which 50% become mature. For C. gariepinus
the percentage of mature fish in each length group was plotted against length for each
sex using data from both breeding seasons (Fig. 10). The average length at maturity for
males is 32 cm and for females 29 cm. The smallest ripe female was 21 cm.
For C. ngamensis the smallest mature male seen during the breeding season was 23 cm
long, the smallest female 18 cm and the majority of fish over 25 cm had active or ripe
gonads.
0 f
0
7btal length (an)

FIG.ll(a).
522 N . G . WILLOUGHBY A N D D . TWEDDLE
e8
I I I I
0 20 30 40
Total length ( c m1
FIG.ll(b).
FIG.1 1 . Relationship between fecundity and total length for (a) C. gariepinus and (b) C. ngamensis.
Fecundity
The fecundity of both species is shown in Fig. 11 as a scatter diagram relating total
number of eggs to the total length of individual fish. The ripe ovaries contained between
600 and 1400 eggs per gram of wet weight for C. gariepinus and 900 to 1800 for C.
ngamensis, with the mean at 1350/g. There was no evidence to suggest that larger specimens
laid larger or smaller eggs than small specimens, or spawned at different times.
Breeding seasons and sites

The breeding seasons shown in Fig. 12 were determined from the numbers of sexually
mature fish taken each month. C. gariepinus below 30 cm and C. ngamensis below 25 cm
which were therefore probably immature were omitted from the calculations. For both
years most breeding occurred during the rainy season between November and March,
with the peak occurring in December, although a few fish were found in breeding condition
in most months. There were considerable differences between the proportions of the C.
gariepinus population which were found in breeding condition at the different sampling
sites. At Chiromo only 5 % of the male fish and 17 % of females were in breeding condition
between November and March, while for this period the figures for Mchacha were 11 %
and 23 % and for Ndombo 25 % and 3 1 % respectively.
Adult C. gariepinus were observed breeding in the shallow water among reed stems in
the vicinity of Chiromo in March 1975, but no larvae or fry of either species were caught
during their first months of life despite several attempts to obtain them.
523
I J
56
J
49
95 58
M
134 78 84
16 69
M
53 24 37
108
F
J
171
D
155
15
ECOLOGY O F T H E CATFISH
N
46
i 23 7
O
89
141
99 138 94 65 25 39 96 156 X
S
A
161
J
46
Males
Females
J
25
M
54
A
98
125
M
199 134 125
I
U
59 46 71
J
D
N
78
I32 69
N = 47
O
IOL
98
-t
30
40
40
A
40t
N=82
P
J
5
4
4
J
4
1
0
M
FIG.13. Seasonal variation in the diet of C. guriepinus.

N. G . WILLOUGHBY A N D D. TWEDDLE
5
5
A
6
8
M
5
8
6
F
61
J
109
D
N
1
7
84
0
67
S
N = 56
A
20
524
TABLEI V
Diet of C. gariepinus. The numbers indicate the number of stomachs in which each category of food item was the dominant constituent
~~~~~~ ~ ~ _ _ _ ~ ~~ ~ ~
Number Fresh Filamentous Plant Dragonfly Other

examined Empty plant algae detritus Mud Humus Chironornids nymphs insects Fish Others
778 293 19 45 109 17 96 20 34 26 110 9

% Diet - 38 3 6 14 2 12 3 4 3 14 1
rn
0
0
TABLE V r
0
Diet of C. ngamensis for theperiod August 1974-July 1976. The numbers indicate the numbers of stomachs in which each category of .food item was the Q
dominant constituent; percentages total 101 "/d due to rounding offigures to the neurest full per cent
Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul Total %total
~~~
No. examined 47 47 42 32 44 42 56 51 51 40 41 48 541 -

Empty 6 17 16 10 12 8 8 24 23 6 17 15 162 30
Plant detritus 3 3 3 6 9 3 2 3 1 1 2 2 38 7
Humus 2 6 2 0 3 5 6 6 6 5 5 4 50 9
Mud n 1 0 0 0 2 0 0 1 0 0 1 5 1
Chironomid larvae 19 4 4 1 5 12 19 9 5 12 10 16 116 21
Beetle larvae 1 0 2 0 2 1 1 0 1 1 1 0 10 2
Dragonfly nymphs 8 11 2 6 1 8 13 5 5 6 1 8 74 14
Mayfly nymphs 2 0 3 1 0 0 0 0 0 6 3 1 16 3
Herniptera 1 2 2 3 1 1 2 0 2 1 0 0 15 3
Molluscs 4 0 1 1 9 0 1 1 2 0 1 0 20 4
Fish 0 3 4 3 1 0 1 1 2 0 0 0 15 3
Others 1 0 3 1 1 2 3 2 3 2 1 1 20 4
526 N . G. WILLOUGHBY A N D D . TWEDDLE
Food and feeding

C. gariepinus
Eight hundred and seventy eight stomachs were examined, of which 725 were from fish
caught in the gill nets, 53 from fish caught by other means at the standard sampling sites,
and the remaining 100 from fish caught on flooded farmland. The results of the analysis
by dominance of the contents of the stomachs taken from fish at the sampling sites are
shown in Table IV. This indicates that the species is omnivorous, stomach contents includ-
ing humus and plant detritus, filamentous algae, insect larvae and fish. Seasonal variations
in some of the more important components of the diet are shown in Fig. 13. The annual
drying of the marsh between September and November can be seen to have a considerable
influence on the diet. A high proportion of the specimens examined at this time had empty
stomachs, and filamentous algae, plant detritus and humus were only rarely consumed.
However fish became important in the diet during these months, probably because they
were being concentrated into a smaller body of water, and were therefore easier for the
C. gariepinus to catch.
There were few significant differences in the diets of specimens taken from the three
sub-stations, although the fish from Chiromo were often found with empty stomachs and
seldom fed on fish, while those from Mchacha seldom fed on filamentous algae. There
were no significant differences between the diets of fish caught at the same site but in
different years.
An analysis which was carried out on the stomachs from 100 specimens caught on
flooded farmland in May 1976 showed, however, that there was a considerable difference
between the diet of these specimens and those from the marsh. The stomachs of 46% of
the former were filled with dragonfly nymphs, 24% with Xenopus tadpoles, 8 % with
notonectid water bugs, and only 5 % were empty. Comparison of these figures with those
shown in Table IV indicates the variation in diet which can occur under different circum-
stances in the same locality.
An attempt was also made to investigate changes in the diet related to the size of fish,
but the only obvious trend was the reduction in the importance of insect larvae among
larger fish. Large specimens were more often found with empty stomachs or with fish in
their stomachs than small specimens, but neither trend was significant and the analysis
was hampered by small numbers of very large and small individuals.
The relationship between the size of C. gariepinus and the size of the fish they consumed
is shown in Fig. 14. The lengths of the prey have been reconstructed to their total lengths
at the time of consumption. The majority of the prey were approximately one eighth the
length of the predator, and very few were larger than one quarter the length. Cichlids were
by far the most common prey, followed by Barbus spp. and other Clarias. One of the
latter was more than half the length of the predator. Clarias were found t o be piscivorous
from a small size, with specimens of 16.5 cm taking cichlid prey.
An index of feeding intensity was obtained by dividing the total weight of food present
in the stomachs examined each month by the total weight of the fish involved. The results
of this analysis, which shows considerably increased food consumption between December
and March, are shown in Fig. 15.
C. ngamensis
The results of the stomach analyses are shown in Table V. Chironomid larvae and
E C O L O G Y O F T H E CATFISH 527
Predotor length ( cm 1
FIG. 14. Relationship between size of predator and size of prey for C. guriepinus.
N = f6 67 84 71 139 61 68 58 65 50 44 45
A S 0 N D J F M .4 M J J
dragonfly nymphs can be seen to be the most important items in the diet. There was little
seasonal variation in the diet, although the fish were consumed mainly during the low
water period of September-November. There were no significant differences in the diets of
fish taken from the different sampling stations.
Discussion
The seasonal variation in the numbers and weights of the Clurius caught in the nets are
probably caused by two factors; changes in the behaviour of the fish and fluctuations in
the area of the marsh, leading to enforced concentration or dispersion of the fish popu-
lations. During December and January the marsh is filling, and the area available for the
528 N . G . WILLOUGHBY A N D D. TWEDDLE
fish is therefore increasing. Maximum catches are taken at this time, indicating that they
are caught as a result of their own activity. It is probable that breeding and increased
feeding activity in this period contribute to these high catches. Low catches between
May and July, while the marsh levels are still high, may be attributed to reduced activity
during the cold season. The water surface temperature is little more than 20°C during
these months, but rises to 30°C in October, and is maintained at this level until March
(R. E. Hastings, pers. comm.). The increases in the catches in September 1974 and
August-September 1975 could be a result of increased activity as the water starts to warm
again, but are more likely to be a result of the drying of the marsh at this time.
Both species were more abundant at Ndombo and Mchacha than at Chiromo, implying
that the marsh and lagoon habitats of the former stations are preferred by both species to
the deeper water of the main river channels sampled by Chiromo nets. C. ngamensis
appears to be more tied to the marsh environment than C. gariepinus and observations
made while electrofishing suggested that this species was relatively uncommon in river
channels. Electrofishing operations also suggested that C. gariepinus prefer to lie on the
substratum, or occasionally in very dense beds of Ceratophyllum, in places where they
have easy access to the air. (These fish die if prevented from reaching the surface in water
which is not well aerated (Donnelly, 1973)). They prefer a cover of vegetation but will also
shelter under mud banks. Consequently large numbers can often be obtained from shallow
weedy swamps, while deeper, more open areas are less productive. However, on occasion
numbers of C.gariepinus were seen surfacing for air in the centre of the main river channel.
The mean weights of specimens taken at the three stations showed considerable variation.
In both species the fish from Chiromo were of a larger average size than those from
Mchacha, which were in turn larger than those from Ndombo. Larger C. gariepinus are
known to prefer deep water (Ratcliffe, 1972; M. N. Bruton, pers. comm.), so it would be
expected that the mean size of fish taken from the main river channel at Chiromo would
be greater than that at the other shallower sites. Also, large numbers of Clurias breed in
the northern part of the marsh around Ndombo, and since the fish do not appear to move
very much it would be expected that the juveniles would make up a larger proportion of
the catch at Ndombo, thereby reducing the mean weight at that site. The juveniles caught
by electrofishing were in shallow water, the smallest being taken in 10 cm of water over
flooded grassland. Since the nets were set in water whose depth was 1.5 m or more, this
may help to explain the scarcity of juveniles in the smaller meshed nets. However, the
scarcity of juveniles in small meshed gill nets is a phenomenon found in many fish (D. H.
Eccles, pers. comm.) and may be a result of smaller fish swimming more slowly and having
less momentum, so being less efficiently caught.
The seasonal fluctuations in the mean weights of both species suggest two inter-related
factors. Juvenile fish are available for capture in the smallest meshed nets at approximately
one year old, and this would depress the mean size of the fish taken during the breeding
season, having less effect as the year progressed. Catches from Chiromo made a more
significant contribution to the overall catch during the colder months of May-June than
at other times of the year, thereby raising the overall mean weight at this time.
C. gariepinus of more than 1 kg were seldom taken in gill nets, despite their presence
in the main river channels. Ratcliffe (1972) reported that the mean size of specimens taken
in experimental nets in Chiromo in 1968 was 4 kg, suggesting that the stocks of large
specimens were not being fully exploited. As none of the local fishermen set nets with mesh
sizes larger than 10.2 cm (4in), this would appear possible, although as experimental
nets with 12.7, 15.3 and 17.9 cm meshes caught only one fish in a year, it is felt that few
fish of such a size now exist in the area.
C. ngamensis has a more limited distribution than C. gariepinus, being apparently
restricted to the marshes and weedy areas of the main river channels, whereas C. gariepinus
is an ubiquitous species, being found in all aquatic habitats including headwaters of
tributaries and pools in dry sandy river beds as well as the main river and marshes.
It is unlikely that significant migrations of C. gariepinus occur, and three out of four
tagged specimens recaptured after more than five months were still within 1 km of their
release points. Fish probably move from the river channels into the nearest freshly flooded
area for breeding purposes at the beginning of the rains. The significant movement of a
single specimen was probably caused by a behavioural change as a result of tagging.
M. N. Bruton (pers. comm.) had very poor returns from tagging studies conducted in
Lake Sibaya, South Africa, but the few recoveries he obtained also showed little movement
from the original release site.
Although conventional techniques for age determination are difficult for tropical species,
several authors have attempted to interpret zones on vertebrae (Bishai & Gideiri, 1965;
Tweddle, 1975), pectoral spines (van der Waal & Schoonbee, 1975), scales (Bruton &
Allanson, 1974) and opercular bones (Lowe, 1952) with some degree of success. Banks,
Holden & McConnell(l965 (unpublished)) in a report on Lake Kainji, Nigeria, considered
10"-11" of latitude to be the limit beyond which rings could be expected to form on
skeletal structures due to changes in metabolic rate as a result of changes in temperature.
The Elephant Marsh lies between 16" 10' and 16" 35' S, and the average monthly tempera-
tures in the marsh differ by 10°C over an annual cycle. It might therefore be expected that
rings would be formed on vertebrae and annual rings were, in fact, formed during the cold
season. At this time food consumption was shown to be considerably less for C. gariepinus,
probably due to reduced metabolic activity at the lower temperatures. Also, at this time
of year water levels in the marsh begin to fall and the fish are being concentrated into a
small area, possibly leading to greater competition for available food. Poor growth and
ring formation are therefore probably due to a combination of these factors. Growth of
C. gariepinus was observed to slow down considerably in the ponds at Kasinthula Fish
Farm, which is supplied by the River Shire, during the colder months (0.Msiska, pers.
comm.). Growth of wild-spawned C. gariepinus juveniles in these ponds is of the same
order as that of marsh fish. The Shire marshes are subject to considerable fluctuations in
water level and area, and a wide range of habitats is available within the marsh environ-
ment, from deep, clear, flowing rivers to extensive areas of shallow swamp susceptible to
deoxygenation and large temperature fluctuations. As a result considerable variation in
growth rates could be expected. Nevertheless, distinct trends are apparent, and reasonable
correlation was obtained for C. gariepinus between growth rates determined by back
calculation of the lengths at which the rings were laid down and the interpretation of
length frequency data for fish up to three years old. For older fish interpretation of length
frequency data is not possible. The positions of the peaks tentatively assumed to represent
3 + year olds underestimate the true modal lengths for that year group, possibly due to
heavy fishing mortality in this size range. The calculated lengths of juveniles during the
cold season agreed well with the few observations on the sizes of juveniles obtained by
electrofishing.
530 N . G . WILLOUGHBY A N D D . T W E D D L E
For C. ngamensis it was not possible to estimate growth rates from length frequency
analysis, and despite extensive electrofishing surveys and trapping experiments very few
juveniles were found, therefore no data are available to confirm the growth rates calculated
from vertebrae. However, as confirmatory evidence is available for C. gariepinus, the two
species occupy similar habitats in the same river system, and the vertebrae have a similar
appearance, it is reasonable to assume that C. ngamensis growth rates calculated from
vertebrae are fairly accurate.
In C. gariepinus growth of both sexes is similar in the first two to three years, after which
the female growth rate appears to slow down, whereas males continue to grow at
approximately 8 cm/year. The majority of large specimens (60 cm +) caught were males.
The period of faster growth coincides with the breeding season, therefore the slower female
growth may be due to more energy being channelled into gonad development in females
than in males. Hastings (1973 (unpublished)) found that one female C. gariepinus had
35 % of its body weight as ovary. It is unlikely that the testes ever exceed 2 % of the body
weight. Van der Waal & Schoonbee (1975) also found that C. gariepinus males grew faster
than females in Transvaal, South Africa.
Male C. ngamensis grow slightly faster than females but the difference is slight and there
were almost as many large females (over 50 cm) as there were males of similar size. The
largest recorded specimen, from the Upper Zambezi, weighed 3.065 kg (Jubb, 1967).
From the length-weight relationship this would have been approximately 68.4 cm in
length, agreeing well with the calculated maximum length of 73.1 cm for males and
69.8 cm for females in the R. Shire.
The average lengths at first maturity, approximately 30 cm for C. gariepinus, 25 cm for
C. ngamensis, agree well with the calculated lengths at the end of the second year. It
appears that for both species most fish therefore breed for the first time at two years old,
with the possibility that some faster growing individuals mature in their first year.
The Shire Valley C. gariepinus first breed at a size similar to that found by Kirk (1967)
for those in L. Chilwa, Malawi, but slightly smaller than that found in L. Sibaya, South
Africa (M. N. Bruton, pers. comm.).
C. gariepinus breeds at the beginning of the rainy season in all the areas for which data
are available (Greenwood, 1955, 1958; Kirk, 1967; Hastings, 1973 (unpublished); van
der Waal, 1974). Van der Waal (1974) considers that some factor which is present in the
run-off water from the first rains is the final stimulant for breeding, and Pott (1969)
suggests that this is an oily substance called “petrichor” which is formed on rock and
soil in hot, dry conditions (Bear & Thomas, 1964). This theory could explain the rapid
development to maturity of specimens at the beginning of the rainy season, but would not
explain the four month breeding season found in the present study. High water levels in
the marshes maintain suitable spawning habitats throughout this period, and it is probable
that the availability of the correct environment is the main cause of the extended breeding
season in this area. The low proportion of CZarias breeding in the Chiromo area is probably
a reflection of the scarcity of flooded !and on which it was possible to set nets. Undoubtedly
reasonable numbers of specimens spawned in areas which were inaccessible to gill
netting.
C. ngamensis also breeds during the rainy season, but the initiation of its breeding
activity seems to be more closely tied to the beginning of the rainy season than in C.
gariepinus. It would appear that the early rains in October 1975 (Fig. 2) caused many
fish to reach sexual maturity unusually early in that year. This was not seen in C. gariepinus
caught over the same period.
In terms of number of eggs per gram of ovary or per gram of body weight, C. ngamensis
lays approximately 50% more eggs than C. gariepinus. However, because of its greater
maximum size the potential fecundity of each female C. gariepinus is greater than that of
C. ngamensis.
Kirk (1967) found regurgitation of stomach contents to be very common in C. gariepinus
caught by gill nets in Lake Chilwa, but it did not appear to occur in the present study as
fish taken in traps had the same frequency of empty stomachs as those caught by gill nets.
The feeding habits of this species may be best described ,as those of a clumsy piscivore.
Greenwood (1958) and Jubb (1967) found that C. gariepims would feed on a variety of
foods, from plankton to fish, while Groenewald (1964) and Kirk (1 967) regarded it primarily
as a predator. Corbet (1961), in a detailed study of the diet of fishes from a variety of
habitats in Lake Victoria and neighbouring waters, stated that C. mossambicus derives
most of its food from Hapluchromis and larger aquatic insects, augmenting this diet with
a wide range of other foods, including plankton, plants and molluscs. Groenewald (1964)
suggested that items other than fish were only consumed as auxiliary or emergency food,
but this was not the case in the present study where insect larvae were of considerable
importance at most times of the year. Also plant detritus and humus were the dominant
items in many stomachs. No experiments were conducted to determine how much of this
ingested matter was actually digestible and therefore a contribution to the nourishment
of the fish. However, it seems unlikely that so many fish would be feeding predominantly
on humus and detritus if these food items were of no nutritional benefit and thus although
the importance of humus and plant detritus in the diet may be overemphasized by the
figures given in Table IV, these items probably do contribute to some extent to the diet
of this species. It is obvious both from the differing accounts in the literature, and from this
study that C. gariepinus is an opportunist in its feeding habits and will consume virtually
anything. Bell-Cross (1974) even records the presence of’ a small crocodile in one C.
gariepinus stomach.
Corbet (1961) stated that after hatching in temporary streams young C. mussambicus
feed mainly on Ostracoda and aquatic insects until they reach a length of about 3 cm,
after which they feed progressively more on fishes. A small sample from torrential reaches
of the Victoria Nile indicated that, despite the different environment, the same ontogenetic
changes in trophic behaviour occur in rivers also.
It is apparent from the literature that no particular group of fish is selected as prey, and
the most common species of suitable size in the area appear to be taken most frequently.
Thus in Lake Victoria C. gariepinus consumes mainly Hoplochrumis spp. (Greenwood,
1958 and Corbet, 1961), in the Transvaal it eats other C. gariepinus (Groenewald, 1964),
and in L. Chilwa it eats Barbus paludinosus Peters (Kirk, 1967).
Jackson (1961) stated that the favourite food of C. ngamensis (under the synonym C .
mellandi Boulenger) was molluscs, but other animal food was also taken. In dams, feeding
habits were reported to be more catholic. In C. ngamensis the vomerine teeth are distinctly
granular and form a very broad band in comparison with the narrow, sometimes inter-
rupted, band of vomerine teeth found in C. gariepinus (see illustrations in Jubb, 1967).
This coarse dentition may be an adaptation to a diet containing a high proportion of
hard-shelled invertebrates.
532 N. G . WILLOUGHBY AND D. TWEDDLE
In the Elephant Marsh system C. ngamensis is a more specialized feeder than C.

gariepinus, and of the fish whose stomachs had contents nearly 60% had been feeding on
insect larvae and nymphs, whereas for C. gariepinus this figure was less than 20%.
Molluscs also occurred in the diet of C. ngamensis, and in 20 of the stomachs examined
were the dominant item. Molluscs were not found in C. gariepinus stomachs, although a
land snail with the shell removed is an excellent angling bait for this species. As in C.
gariepinus, stomachs of C. ngamensis were found to contain a wide variety of potential
food items, including plant detritus, molluscs, insect larvae and fish.
It is difficult to assess the causes of the seasonal variations in the diet of C. ngamensis.
The most obvious is that the chironomid larvae, which make up the bulk of its diet, are
relatively uncommon during the dry marsh period of September-November. More fish
were consumed during this period than at any other time of year, a phenomenon also
found for C. gariepinus, although fish never formed a very important part of the diet of
C. ngamensis. This species must be classed as an omnivore, but appears to prefer insect
larvae when they are available.
Summary
Clarias gariepinus and C. ngamensis together make up approximately 70% of the
traditional fishery catch of 12-14,OOO tons per year from the River Shire marshes. Research
into their ecology was conducted using graded fleets of gillnets at three stations around
the Elephant Marsh. Peak catches occurred in December-January and in August-
September. The former peak was attributed to increased activity during the breeding
season, and the latter to falling water levels which allowed the fish to be captured more
easily as they were forced off the floodplain into the main river channels.
C. ngamensis was found to be restricted mainly to the marsh and lagoon systems,
whereas C. gariepinus is an ubiquitous species, being found in all aquatic habitats including
headwaters of tributaries and pools in dry sandy river beds as well as the main river and
marshes.
Movement of C. gariepinus within the marsh system was investigated by tagging.
Seventeen percent of the marked fish were recaptured, and few had moved significant
distances from the original point of capture.
The growth rates of both species were determined by back calculation from the hyaline
rings formed on the vertebrae during the cold season, May-August. Length frequency
analysis, tagging studies and electrofishing surveys provided additional information on
growth of C. gariepinus. This species had a faster growth rate than C. ngamensis and
attained a much larger size, the largest C. gariepinus caught being a 100 cm male and the
largest C. ngamensis a 55 cm male. In both species males grew faster than females, the
difference between the sexes being most marked in C. gariepinus.
The sex ratio was significantly different from unity, with, in C. gariepinus 9 % more,
and in C. ngamensis 20% more males than females being caught.
Breeding in both species occurred during the rainy season, from November to March,
with the peak in December to January. Lengths at maturity agreed well with calculated
lengths at the end of the second year in both species. C . gariepinus ovaries contained up to
160,000 eggs, with about 900 eggs/g wet weight, whereas C. ngamensis ovaries contained
up to 50,000 eggs with an average of 1350 eggs/g wet weight.
ECOLOGY O F T H E CATFISH 533
C. gariepinus was omnivorous and opportunist in its feeding habits and plant matter,
humus, insect larvae, tadpoles and fish were all important components of the diet at times.
The piscivorous role of C. gariepinus was only significant when the marsh water level was
low during October and November. Most of the prey were small cichlids of about one
eighth to one quarter the length of the predator, although Barbus and other Clarius were
also taken. Feeding was most intensive between December and March. C. ngamensis,
although also omnivorous, was a more specialized feeder, with a preference for insect
larvae.
We would like to thank Mr D. H. Eccles and Dr D. S. C . Lewis for useful comments and
criticisms and Ms S . C. Tuohy for typing the manuscript.
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534 N . G . WILLOUGHBY A N D D. TWEDDLE
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Journal of Zoology - December 1978 - Willoughby - The Ecology of The Catfish Clarias Gariepinus and Clarias Ngamensis in

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Journal of Zoology - December 1978 - Willoughby - The Ecology of The Catfish Clarias Gariepinus and Clarias Ngamensis in

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J . Zool., Lond.

(1978) 186, 507-534

The ecology of the catfish Clarias gariepinus and Clarias ngamensis

Description of study area

1974 ' I975

Age and growth

b t a l number and weight

lndividuol weigMs (Smoothed means)

1974 1975 I976

Migra tion-C. gaviepinus

Time belween logging and recapture (days)

Age and growth

Length d fish (crn)

given at right of figure.

Growth rates calculated from vertebrae

Jan '75 July

Growth of tagged fish-C. gariepinus

C . gariepinus: log W = 3.136 log L-2.3396

Length when Length when

292 28.0 34.0 6.0

7btal length (an)

Breeding seasons and sites

FIG.13. Seasonal variation in the diet of C. guriepinus.

Number Fresh Filamentous Plant Dragonfly Other

778 293 19 45 109 17 96 20 34 26 110 9

No. examined 47 47 42 32 44 42 56 51 51 40 41 48 541 -

Food and feeding

In the Elephant Marsh system C. ngamensis is a more specialized feeder than C.

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