Acta Tropica, 61(1996)191-199 191

© 1996 Elsevier Science B.V All rights reserved 0001-706X/96/$15.00

ACTROP 00527

Food selection behaviour of potential biological

agents to control intermediate host snails of
schistosomiasis: Sargochromis codringtoni and
Tilapia rendalli
M o s e s J. C h i m b a r i a,,, Jerikias N d a m b a b, H e n r y M a d s e n c
a De Beers Research Laboratory, P. 0. Box 197, Chiredzi, Zimbabwe
b Blair Research Laboratory, P. 0. Box CY573, Causeway, Harare, Zimbabwe
c Danish Bilharziasis Laboratory, Jaegersborg Alle 1D, DK-2920, Charlottenlund, Denmark
(Received 18 July 1995; revision 27 November 1995; accepted 27 November 1995)

The food selection behaviour of two fish species indigenous to Zimbabwe (Tilapia rendalli and
Sargochromis codringtoni) was studied under laboratory conditions with a view to considering them as
biological agents for snail control. Six glass aquaria were set up and divided into two sets each with two
aquaria. One S. codringtoni was introduced into each aquarium of the first set while one T. rendalli was
introduced into each aquarium of the other set. In one of the aquaria of each set the fish were supplied
with snails and trout food. Snails and weeds were provided, but trout food was excluded in the second
aquarium of each set while the third aquarium for each set was provided with trout food, snails and
weeds. A fourth aquarium with weeds and snails but no fish was set up as a control. Data collected over
9 weeks showed that T. rendalli was primarily herbivorous while S. codringtoni was shown to be
malacophagous. Presence of trout food made no difference in the snail-eating habit of S. codringtoni or
the weed-eating behaviour of T. rendalli. However, trout food seemed to be a good protein supplement
to T. rendalli as the fish with access to trout food gained more weight and length.

Key words: Tilapia rendalli; Sargochrornis codringtoni; Snail control; Schistosomiasis; Biological control;
Laboratory

1. Introduction

Snail control is a major component of any successful schistosomiasis control pro-

gramme (Gundersen et al., 1990). Currently, the most widely used method for
control of snails is application of toxic chemicals in snail habitats. The recommended
chemical is Bayluscide which has been demonstrated by several control programs to
be very effective in killing snails (Evans, 1983; Gundersen et al., 1990).
Unfortunately, the cost of Bayluscide (US$78 300/ton if purchased through Bayer-
Zimbabwe) has increased drastically in the past few years to such an extent that it
is no longer affordable to the third world countries where schistosomiasis is endemic.
In Zimbabwe, use of Bayluscide is confined to large irrigation schemes and tourist

* Corresponding author. TeL: (+263-131) 2351;fax." (+263-131) 2209.

SSDI 0 0 0 1 - 7 0 6 X ( 9 5 ) 0 0 1 4 4 - 1
192

recreation centers that can afford the expenses involved. This situation is not satisfac-
tory since the majority of the exposed population lives in the communal, resettlement
and small-scale irrigation areas. There is therefore justification for studies that seek
to introduce alternative affordable snail control methods.
Theoretically, biological methods for controlling undesirable disease-causing
agents are environmentally sound. However, there are several practical implications
that have to be taken into account before such methods can be adopted. A successful
biological agent must preferably be indigenous and should prefer a habitat that
coincides with the habitat of the target agent. Such an agent should be one that
prefers the target agent even if alternative food is available. Furthermore, there must
be adequate knowledge of the long-term effects on the environment of using biologi-
cal control agents.
In view of the above considerations, the present study was conducted to study
the food selection behaviour of fish indigenous to Zimbabwe that have a potential
for controlling intermediate host snails for schistosomiasis. Tilapia rendalli is a
primarily herbivorous fish that was once used in the Lowveld of Zimbabwe to
control weeds in irrigation channels (Lingen et al., 1960). This fish was shortlisted
as a candidate for snail control on the rationale that removal of weeds from snail
habitats reduces the amount of food available to the snails and may subject the
snails to natural predators. Sargochromis codringtoni is known to be a snail eater
(Bell-Cross and Minshull, 1988), but the extent to which it could be used for
controlling snails is not known.
No food selection behaviour studies similar to the present one have been conducted
on either S. codringtoni or T. rendalli. One study conducted on the former fish
focused on gathering evidence of the malacophagous behaviour of the fish and on
establishing the stocking densities of the fish that would result in the elimination of
snails in an ideal situation where snails are the most abundant food (Larson and
King, 1994).

2. Materials and methods

Seven glass aquaria measuring 100 × 60 × 60 cm were constructed. A sand substrate

with a depth of 8 cm at one end (length wise) and 4 cm at the other end was
provided in each aquarium. Twenty aquatic plants (Vallisneri sp.) were planted in
each aquarium at similar positions. The aquaria were filled with natural filtered
pond water and left to stand for 1 week. Twenty 5-week-old snails of each of the
species Bulinus globosus, Biomphalaria pfeifferi (both intermediate host snails for
schistosomiasis) and Bulinus tropicus (a non-intermediate host) were introduced into
each aquarium. The snails were fed on sun-dried lettuce for 4 weeks after which the
following treatments were made to each aquarium (AQ):
AQI: all aquatic plants were removed and 2 female T. rendalli were introduced
AQ2: two female T. rendalli were introduced
AQ3: two female T. rendalli were introduced
AQ4: no fish were introduced (control)
AQ5: all aquatic plants were removed and 2 male S. codringtoni were introduced
AQ6: two male S. codringtoni were introduced
AQ7: two male S. codringtoni were introduced
 193

It was not possible to maintain the same sex for both species of fish as no matching
sizes (weight and length) were available. The sizes of the experimental fish ranged
between 13 and 17 cm in length and between 50 and 60 g in weight. The number of
fish in each aquarium was reduced to one after 1 week. The fish were removed
because S. codringtoni showed cannibalistic tendencies when snails were in short
supply. It was therefore decided to remove the weaker fish in the case of S. codringtoni
and in the case of T. rendalli removal was such that the remaining fish in each
aquarium would be similar to fish in the other aquaria in terms of length and weight.
Feeding routines for fish and snails in each aquarium were controlled: 0.5 g of
lettuce was added to each aquarium on Monday and Wednesdays only; 1 g of
commercial trout food was added to aquaria 1, 3, 5 and 7 every day; 100 g of wet
Najas pectinata was added to aquaria 2, 3, 4, 6 and 7 every Monday. Every week
equal numbers of snails by species were introduced into each aquarium. The number
of snails added per week varied slightly from one week to another but a minimum
of 20 for each of the three species of snails was added in each aquarium.
Every Friday throughout the 9-week study period, 5 traps were immersed in each
aquarium to sample for snails. The traps were four cornered plastic bottles measuring
4 x 4 cm with a mouth diameter of 3 cm. One surface of the bottle was cut out and
replaced by a mesh cloth with an opening at the center (diameter 1.5 cm) to allow
snails to crawl into the trap. A piece of thread was tied to the bottle neck using the
screw cap and the other end was attached to a kaylight float to allow the trap to
move freely in the water. Dried lettuce (1.5 g) was put into each trap as an attractant
for snails and a small weight (stone) was put inside the trap to suspend the trap in
the water. Once the traps were introduced, all feeding either with trout food or
lettuce was suspended.
The traps were removed from the aquaria every Monday and the snails in each
trap were classified according to species and size and counted. After counting, the
snails were returned to their respective aquaria. The amount of N. pectinata remain-
ing in each aquarium where it had been introduced was weighed and returned to
the respective aquaria afterwards.
Routine observations were made for 1 h every day to take note of any peculiar
behaviour by the fish, particularly their feeding habits.

3. Results

3.1. Impact offish on snails

Fig. 1 shows the results of weekly snail sampling made in each aquarium for 9
weeks. A marked decline in snail numbers from sampling performed in week 1 (just
before introduction of fish) to week 2 was observed in all the aquaria except in the
control aquarium (AQ 4) where the reduction was minimal. While very few snails
were sampled in aquaria containing T. rendalli (AQ 1-3), no snails were sampled
in aqua~a containing S. codringtoni (AQ 5-7) in week 2 and the rest of the sampling
period (9 weeks). In aquaria containing T. rendalli the numbers of snails detected
fluctuated, with more snails generally being detected in the aquarium where trout
food was added (AQ 3). The control aquarium (AQ 4) maintained higher numbers
of snails than any aquarium, reaching a peak of 80 snails in week 7.
194

100

SO

AQUARIUM 1
-4- AQUARIUM 2

~ - AQUARIUM 3
4 - AQUARIUM 4
40 ¸ -x- AQUARIUM 6
W
-*- AQUARIUM 6
-'- AQUARIUM 7
20

0
2 3 4 8 6 7 8 9
WEEK

Fig. 1. Snail counts before introduction of fish (week 1) and after introduction of fish (weeks 2-9). Key:
AQ 1 = T. rendalli + snails + trout food + lettuce; AQ 2 = T. rendalli + snails + weeds + lettuce;
AQ 3 = T. rendalli + snails + weeds + trout food + lettuce; AQ 4 = snails + weeds + lettuce; AQ
5 = S. codringtoni + snails + trout food + lettuce; AQ 6 = S. codringtoni + snails + weeds + lettuce;
AQ 7 = S. codringtoni + snails + weeds + trout food + lettuce.

The results obtained when all the aquaria had water drained and all snails counted
by species are shown in Fig. 2. No live snails or shells of snails were found in aquaria
that contained S. codringtoni (AQ 5-7). Bulinus globosus were found to be predomi-
nant in all the aquaria where snails were present followed by Bulinus tropicus.
Biomphalaria pfeifferi was found in small numbers. Contrary to observations made
during the sampling period, there were more snails found in aquarium 3 which
contained T. rendalli with trout food than in the control aquarium. The number of
snail shells counted in the aquaria still indicated that there were more snails in
aquarium number 3.

3.2. Impact offish on weeds

Fig. 3 shows the total amount of N. pectinata (wet weight in g) present in each
aquarium at the time of snail sampling. There was a marked difference in amount
of plant material between aquaria containing the different fish species, with aquaria
containing S. codringtoni having more plant material.The control aquarium had the
largest quantity of plant material throughout the sampling period.
Final quantification of plant material in each aquarium at the end of the experi-
ment showed that aquarium 4 (control) had the largest quantity of plant material,
followed by the aquarium with S. codringtoni plus trout food (AQ 7) and lastly
aquarium 6 with S. eodringtoni but without trout food (see Fig. 4). There was very
little plant material left in aquaria where T. rendalli was present. Comparatively
 195

350

300-

u) 200 -
[ ] B.globosus
[ ] B.troplcus
~150-
• B.pfelfferl

0 ~ I I I I

1 2 3 4 5 6 7
AQUARIA NUMBER

Fig. 2. Total number of snails recovered in each aquarium at the end of the experiment.

350

!
AQUARIUM 2
AQUARIUM $
-'- AQUARIUM 4
-*- AQUARIUM 8

|,-t / -,k-AQUARIUM 7

0 1 2 3 4 IS 6 7 8 9
WEEK

Nots: aquaria 1 shd 5 are Iw~luded as the weeclawere remowd at the beginning of the exp~m,,m

Fig. 3. Amount of vegetation (wet weight in g) in each aquarium before (week 0-1) and after (week 2-9)
introduction of fish.
196

35O

30O

250
A

I"

ilS0 -

100-

5O

0 i
2 3 4 6 7
AQUARIA NUMBER

Notll: a q u a r i a 1 a n d 8 M e mxt:klded ,u~ t h e weeds were removed l i t ~ e b e g i n n i n g o f the e~perlrnent

Fig. 4. Amount of vegetation (wet weight in g) recovered in each aquarium at the end of the experiment.

there was more plant material in the T. rendalli aquarium where trout food was
added than in the other aquaria containing the same fish.
Table 1 shows the data on fish used in the experiments. All the fish gained weight
and length during the study period. Generally, T. rendalli gained more weight than
S. codringtoni. Among T. rendalli, the fish in the aquarium where trout food was
added gained much more weight and length than the other fish, whilst among S.
codringtoni the fish in the aquarium where no trout food was added gained more
weight and length than the other fish.

3.3. Routine observations

On many occasions S. codringtoni were observed searching for snails, locating

them and sucking them before crushing the shell to gain access to the soft parts.

Table 1
Experimental fish growth data

Aquarium Fish Old wt. (g) New wt. (g) Old length (cm) New length (cm)

1 Tilapia 80 92.6 17 17.5

2 Tilapia 60.7 64.2 15 15.7
3 Tilapia 55 79.1 14 16
4 no fish no fish no fish no fish no fish
5 Sargo 69.7 77.7 16 16.2
6 Sargo 70 80.2 15.5 17.5
7 Sargo 51.4 54.4 14.5 14.5
 197

Fragmented small pieces of shells were spat out in the case of medium to large snails
(shell height > 5 mm) while snails of shell height less than 5 mm were swallowed
without being crushed. Snails with shell heights greater than 10 mm were crushed
after several attempts.
T. rendalli was rarely observed feeding on weeds during the day. However, it was
noticeable every morning that a substantial amount of weeds had been consumed
either overnight or at times when the fish were not under observation. T. rendalli in
the aquarium without weeds (AQ I ) burrowed in the sand continuously, apparently
searching for food. This resulted in the water becoming very turbid throughout the
experiment. It was also observed that T. rendalli ate some of the lettuce in the traps.

4. Discussion

The malacophagous behaviour of S. codringtoni documented by other

Zimbabwean authors (Mitchell and Gahamadze, 1976; Bell-Cross and Minshull,
1988) was confirmed in this study. The absence of snails or shells of snails at the
end of the experiment, in aquaria where S. codrington was present, suggests that the
snails were eaten by the fish. The feeding habits noted during routine observations
seem to be characteristic of Cichlidae fish. Chiotha et al. (1994) also noted the
sucking of snails and crushing of shells. The same authors observed that the fish
(Trematocranas placodon) they were working on had problems attacking snails
clinging to glass walls or on weeds. In another study Acra et al. (1986) noted that
Gambusia affins could not eat snails with shell heights greater than 6 mm. In the
present study all snails were eventually eaten regardless of size, and snails on glass
walls or weeds were no exception.
The observation that snails introduced into the aquarium where trout food was
added and the aquarium where trout food was not added were finished at the same
time in the case of S. codringtoni suggests a preference for snails as opposed to trout
food by the fish. The importance of weeds as a refuge against predators by snails
(Coates, 1984; Thomas and Daldorph, 1991 ) was apparent as the fish in the aquarium
without weeds finished snails much faster than the other fish. The high affinity for
snails exhibited by S. codringtoni cannot be overemphasized as a single fish ate over
600 snails within 9 weeks. More snails could have possibly been eaten had they been
supplied.
The presence of many snails in the aquaria with T. rendalli at the end of the
experiment demonstrated that this fish is not malacophagous. Presence of more
snails in aquarium 3 with T. rendalli than in the control aquarium (AQ 4) further
emphasized this point. The fish in aquarium 1 where there were no weeds, however,
appears to have preyed upon small snails as few small snails were recovered from
this aquarium. It is possible that the fish were forced to eat small snails because of
shortage of food. Alternatively, the small snails may have been selected against by
the unfavourable turbid conditions in that aquarium (El Gindy, 1957). The absence
of weeds where eggs are usually deposited could also explain the absence of small
snails.
Consumption of about 900 g of N. pectinata in 9 weeks indicated that T. rendalli
is primarily herbivorous. This reaffirms observations made by Coates (1984) and
justifies the use of the fish for weed control (Lingen et al., 1960). Contrary to
observations made in T. rendalli, S. codringtoni was not found to be herbivorous.
198

The amount of N. pectinata that could not be accounted for at the end of the
experiment in aquaria with S. codringtoni was negligible.
The attainment of more weight and length by T. rendalli when compared with S.
codringtoni is not surprising considering the amount of biomass consumed by T.
rendalli. The differences in growth for fish (both species) in aquaria where trout
food was added and fish in aquaria without trout food could have dietary implica-
tions. For T. rendalli, fish in aquaria with trout food gained more weight and length
possibly because of the supplementary protein-rich commercial food. For S. codring-
toni, the fish in the aquarium without trout food gained more weight and length,
suggesting that the added trout food did not give the fish with trout food any dietary
advantage.
Overally, this study has shown that T. rendalli has great potential as a weed
controller. This is important from an agricultural point of view (water saving and
infrastructure maintenance) as well as in controlling snails that transmit schistosomi-
asis through removal of weeds (Daldorph and Thomas, 1991). The potential of S.
codringtoni as a direct control for snail populations was demonstrated. The preference
of T. rendalli for weeds and that of S. codringtoni for snails in the presence of
alternative food for both cases has important practical implications in the fishes'
use as biocontrol agents. Very often biocontrol agents that show great potential in
the laboratory fail in the field because of many unchecked variables. More experi-
ments in a seminatural environment therefore need to be conducted on T. rendalli
and S. codringtoni in accordance with WHO specifications before field trials on snail
control can be done (World Health Organization, 1984). The possibility of using
both fish in the same habitat for best results needs to be investigated. The cannibalis-
tic tendencies exhibited by S. codringtoni should, however, be investigated before
both fish are put in the same habitat, as this may lead to elimination of T. rendalli
if snails become scarce and therefore result in re-establishment of the weeds and
consequently of snails.

Acknowledgements

We thank the Lake Kariba Fisheries Research Institute and the University Lake
Kariba Research Station for their logistical support in the catching of S. codringtoni
in Lake Kariba; and Mtirikwi National Parks station for supplying us with T.
rendalli. We are grateful to O. Rubaba, D. Ndlela and M. Viriri for assisting in the
technical work and particularly B. Ndlela for assisting in the catching of S. codring-
toni. This project was supported by DANIDA through the Danish Bilharziasis
Laboratory. Our gratitude goes to the permanent secretary in the Ministry of Health
and Child Welfare for granting us permission to publish this article and to Dr.
Chandiwana, director of Blair Research Institute, for his encouragement and
support.

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 199

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