Aquaculture, 30 (1983) 335-351 335

Elsevier Scientific Publishing Company, Amsterdam - Printed in The Netherlands

A COMPARISON OF THE BIOLOGICAL CHARACTERISTICS OF

SAROTHERODON NILOTICUS (L.) WITH THOSE OF S. AUREUS
(STEINDACHNER) AND OTHER TILAPIA OF THE DELTA AND LOWER
NILE

A.I. PAYNE and RI. COLLINSON*

Biology Department, Couentry (Lanchester) Polytechnic, Coventry (Great Britain)
*Research and Development, Atkins and Partners, Epsom, Surrey (Great Britain)
(Accepted 19 January 1982)

ABSTRACT

Payne, A.I. and Collinson, R.I., 1983. A comparison of the biological characteristics of
Sarotherodon niloticus (L.) with those of S. aureus (Steindachner) and other tilapia of
the delta and lower Nile. Aquaculture, 30: 335-351.

The tilapia species occurring in the lower Nile are Sarotherodon niloficus, S. aureus,
S. galiaeus and Tilapia tittii. The distinguishing characteristics between the previously con-
fused S. niioticus and S. aureus are summarised; from these there is no evidence of hybri-
dization of the two species in natural populations in Egypt. The scales are used to estimate
the growth rates of tilapia species in two coastal lakes and in both, S. niloticus grows
faster than S. aureus after the first year. The possible factors causing the growth checks on
the scales are considered.
The spawning season of S. niloticus appears to attain a discrete peak in April-May,
whilst the spawning season of S. aureus extends from May to September with at least two
actual spawnings within this period. Natural spawning cycles are compared with those
observed in fish ponds.
A more extended spawning period of S. aureus may explain the reduced growth rate of
the species after the first year. The spawning cycle of all species coincides with the tem-
perature regime of the water bodies. The fecundities of S. aureus and S. niloficus are
similar and are described by log F = log 1.33 + 2.23 log L, which suggests that small fishes
produce more eggs per g body weight than large.
The salinity tolerance of the Nile tilapia can be ranked as T. zillii > S. galilaeus > S.
aureus > S. niloticus. Evidence on chronic and acute effects of salinity are reviewed and
upper estimates for salinities giving unimpeded growth are deduced as being, T. tillii 29’/,, ,
S. galilaeus 15-20°/,0, S. areus 10-150/,, and S. niloticus 5-loo/,,.

INTRODUCTION

Surotherodon niloticus (L) and S. aureus (Steindachner) have been used

extensively for fish culture in both Africa and the Middle East as well as in
various other parts of the world to which they have been introduced. The two
species were only distinguished relatively recently (Fishelson, 1962; Trewavas,

0044-8486/83/0000-0000/$03.00 o 1983 Elsevier Scientific Publishing Company

336

1965) and there has been some confusion by workers both before and since
with respect to the precise species they have been dealing with. In Israel, for
example, the status of S. niloticus is questionable in that the natural distribu-
tion is probably restricted to a few coastal rivers, principally the Yarkon,
rather than extending throughout the country as was previously thought
(Fishelson, 1966). Here the situation is made less clear because under certain
conditions the two will hybridize, often to produce a skewed sex ratio, which
can be beneficial if controlled but can add to the complexities if not strictly
regulated (Pruginin et al., 1975; Mires, 1977). The recorded range of S. aureus
extends to the “soudanian” regions of West Africa including the middle Niger
and the Chad Basin, the lower Nile and most of Israel (Trewavas, 1965).
To characterise these two species more fully the present intention was to
examine some of their more important biological features, particularly with
a view to including their use for fish culture since the work was carried out
in connection with a resource survey for a major fish farm development in
Egypt. In addition to the basic processes of growth and reproduction, the
influence of the most significant environmental variable within the Nile delta,
salinity, was also examined. Beyond the optimum range for a species, salinity
can inhibit the growth of fishes well before the upper lethal limit is obtained
(Payne, 1979).
Within the Nile delta, S. niloticus and S. aureus co-exist with S. galileus
(L) and Tilapiu zillii Gervais and since they have also frequently been used
for culture some comparative data were also collected on these species.

METHODS

Fish samples were obtained from fishermen’s catches from the landing
ground by the desert road on L. Maryut and also landing areas at Dumietta
and Port Said on L. Manzalla during April and August. The fish were examined
for distinguishing meristic features, reproductive state and measured for
standard length. A number of scales, including a key scale, were taken from
each individual and mounted directly on a microscope slide where the scale
rings and total radius were measured with an eye-piece micrometer.
A key scale analysis was carried out using the scale above the anus in the
third row on the flank. Scales from this region were used for examination as
experience suggested that there were consistently fewer replacement scales
in this area and rings were generally clearer. The average length when the
various rings were layed down can be backcalculated from the following
formula (Tesch, 1968):

L?l _c =-
sn(L-c)
s

where L, = length of fish when ring “n” was formed; L = length of fish at
capture; S, = radius of check “n”; S = total scale radius; c = intercept on
length axis.
 331

To estimate fecundity the ovaries were first kept in formalin for several
days to render the eggs more durable and then the number of mature eggs in
one ovary was counted and then doubled to give the total number produced
by the fish.
To determine salinity an Electronic Switchgear Conductivity meter was
used which had first been calibrated for salinity against a serial dilution of
seawater.

IDENTIFICATION AND DISTRIBUTION OF EGYPTIAN TILAPIA

A summary of the field characters of the four tilapia species in Egypt has
been given by Payne and Collinson (1979). The most reliable differences be-
tween S. niloticus and S. aureus, however, are the higher dorsal spine count
in the former and also that the lappets of the dorsal fin are always dark, never
red or pale. In S. niloticus from Egypt 83.3% had 17 dorsal spines and 10.4%
had 16 whilst for S. aureus the comparable proportions were 10.3% and 81%.
The two species are, however, readily distinguishable by colour and other
factors in the field.
All four species were found in the shallow, highly productive northern
coastal lakes of Maryut, Idku and Manzalla, in the various canals and drains
of the delta irrigation system ,and also in Government fish farms of the delta.
It has been indicated that S. aureus is not found in the Nile more than a
few kilometers south of Cairo (Trewavas, 1965) and observations 100 km
upstream of Cairo in the present survey concur with this. Similarly, S. aureus
was not found in the catches from Lake Nasser where S. galilaeus has super-
ceded S. niloticus as the predominant species with T. zillii being of subsidiary
importance. It appears, therefore, that S. aureus is confined to the Nile delta.

AGE AND GROWTH

Use of the scales to determine age

Scale rings have been used bpreviously to determine age and growth of
tilapia in Egypt (Jensen, 1957; El Zarka et al., 1970) and in Israel (Ben-Tuvia,
1959). Since, at these latitudes, there is a distinct warm and cool season it
was assumed that the rings were winter rings although Ben-Tuvia (1959) did
suggest, following Holden (1955), that they may have been associated with
spawning. The data in Table I provides some evidence that ring formation is
associated with spawning. S. niloticus appears to have a peak in ring forma-
tion associated with a peak of spawning activity in April/May whilst S. aureus
shows a build up of ring formation through the summer and seems to have a
more protracted spawning season. Jensen (1957) also found the scale rings
of S. niloticus in L. Maryut were formed between March and May.
Many fish examined during April showed severe erosion at the edge of the
scales and the ring itself, when formed, showed severe disruption and repair
338

TABLE I

Proportion of adult fish which are ripe and proportion with ring just formed at the edge
of the scale between February and August 1979 for Sarotherodon aureus and S. niloticus

Site Date Species Ring at Number Number

scale edge (%) ripe % of fish

Various February Sn + Sa 0 0 56
L. Maryut 1414179 Sa 0 87.5 16
Sn 46.6 66.6 15
L. Manzalla 7/5/79 Sa 19 85.7 21
Sn 20 20 5
L. Manzalla 16/S/79 Sa 52.2 82.6 23
Sn 4.5 13.6 22

Sa = S. aureus; Sn = S. niloticus.

Fig. 1. A scale from Sarotherodon aureus showing the double ring of the erosion type at
the outer locus which is probably associated with the spawning cycle. Specimen 14.7 cm
SL, taken from Lake Manzalla.
 339

Fig. 2. A scale from Sarotherodon aureus which demonstrates the contrast between the
faint inner “juvenile” mark and the two outer checks of the erosion type which also show
traces of being double. Specimen 14.3 cm SL, taken from Lake Manzalla.

of the circuli (Figs. 1 and 2) which is often associated with spawning rings
(Cragge-Hine and Jones, 1969; Payne, 1976).
Similar rings have also been found associated with the spawning of T.
guineensis and S. melanothron (AI. Payne, unpublished data) and in
the 2+ age group of the cichlid Sergochromis coderingtoni (Holcik, 1974).
The sequence of events appears to be that as the gonads mature the increased
demand by the gonads for calcium causes resorption from the scales which
are then repaired when maturation is complete leaving what is virtually a scar
on the scale (Garrod and Newell, 1958; Payne, 1976). In temperate fish, al-
though spawning rings are sometimes found, the more typical ring is charac-
terised not by a disruption of the circuli but by “cutting over” which seems
to result not from a transition from cold to warm temperatures tut a change
340

from slow winter to rapid summer growth due to rapidly increasing food
supplies, and consequently the rings do not appear at the edge of the scale
until May-July (Jones, 1953; Cragge-Hine and Jones, 1969). In the main body
of L. Maryut the increase in phytoplankton productivity begins between
April and May and reaches a peak in July (Aleem and Samaan, 1969) and
thus the main transition of food availability, like gonad maturation, coincides
with ring formation at the edge of the scale, at least in S. niloticus.
The fact that size at first maturity exceeded the size when the first annual
ring is laid down in S. niloticus (Table IV) may emphasize the complementary
effect of food availability on spawning in this species although it is also pos-
sible that the true size at first maturity was not found, since in the Nile at
Khartoum the minimum sizes for males and females is 11.4 and 14.3 cm
(Babiker and Ibrahim, 1979) which coincides with the size at which the first
“erosion” type ring is laid down in lower Egypt.
In a proportion of the fish examined a second type of ring was found. This
was invariably the innermost ring and its structure closely resembled the “cutt-
ing over” pattern typical of temperate fish rather than the “eroded” structure
mentioned above and it was always formed when the fish was small, between
2-6 cm. This was taken to represent fish which were spawned late on in the
season, around August or September, which consequently were well below
the adult size the following spring and would, therefore, not primarily be
influenced by the spawning cycle at this time. This type of ring could be
caused by increasing food supply as mentioned above or by a “trial run” of
the endocrine system controlling reproduction. In L. Maryut 31% of S. nilo-
ticus and 35% of S. aureus possessed a first ring of this type. El Zarka et al.,
(1970) commented upon the occurrence of this relatively faint inner ring but
did not give any explanation or say how it was treated in the analysis. A
similar “juvenile” ring was found in fishes from Lake Kariba where for exam-
ple it was seen in 61% of S. mortimeri (Krupka, 1974) and 91% of Surgo-
chromis codringtoni (Holcik, 1974) examined as well as in some noncichlids
such as Hydrocynus uittutus (Balon, 1971).

TABLE II

A comparison of annual size estimates (cm) derived from both anal and shoulder scales
from Sarotherodon aureus and S. niloticus, Lake Manzalla, Egypt

Year S. aureus S. niloticus

Anal Shoulder Anal Shoulder

1 9.8 9.0 8.2 7.7

2 14.4 14.0 16.8 16.8
3 16.5 16.4

Number
sampled 26 19
 341

In S. aureus traces of two rings at a single location could often be clearly

discerned which may indicate at least two spawnings in a season (Fig. 1).
Table I seems to suggest that the earliest spawning of S. uureus is not accom-
panied by the formation of a ring. Holden (1955) suggests that if talapia
maintain their condition, spawning marks are nor formed. Consequently, in
the case of S. aureus, only the energy strain imposed by subsequent spawnings
during the year may induce the formation of checks on the scales.
The relationship between total body length and scale diameter (cm) was
found to be:
L = 0.2673 + 1.6
where L = length (cm) and S = scale diameter (arbitrary micrometer units).
A comparison was carried out with scales from the third dorsal row, on the
shoulder, just behind the operculum (Table II) and this demonstrated that
differences were generally small. At the youngest ages the rings on the
shoulder scales tended to be formed at a slightly smaller size and in some
individuals it could be seen that whilst rings on the anal scales were still being
formed at the edge those from the shoulder had been formed and a small
subsequent amount of scale growth had taken place. Repair of the scale fol-
lowing erosion of the scale edge apparently, therefore, can occur slightly
earlier in the shoulder than in the anal scales.

Growth rates

The mean lengths at the end of each year for the four tilapia species from
a variety of locations both from the present work and from that of others are
shown in Table III. All measurements were initially taken as standard lengths
(SL) but were corrected to total lengths (TL) after the relationship between
the two from a sample of 40 fish was found to be:
TL = 1.33 SL’“’
The estimate of growth rate for S. niloticus in L. Maryut although based on
small numbers, compares closely with that obtained in previous estimates
from the same lake (Jensen, 1957; El Zarka et al., 1970).
Although one of the commonest tilapia of the delta, S. aureus has rarely
been distinguished from other species. Jensen (1957) in his study of growth
of tilapia species in Lake Maryut makes no mention of it and, therefore, pre-
sumably may have confused it with S. niloticus or S. galilueus in his estimates.
El Zarka et al. (1970) did distinguish it in that they noted that whilst “Tilupia
nilotica Linn” was common in the lake the most frequent was a type referred
to as “2’. nilotica (variety x)“. This must have been S. aureus and, moreover,
they excluded it from their estimates of growth for S. niloticus. S. aureus is
important because it has been used very successfully for culture in Israel and
has generally been founded to be the most satisfactory species to use in that
area where the environmental conditions are very similar to those in Egypt.
342

TABLE III

Mean annual lengths of tilapia species from various localities in lower Egypt (n = no. fish
sampled)

Location and Mean annual length (cm and standard deviation (u )

species
1 01 2 02 3 03 4 *4 n

L. MANZALLA
S. niloticus 9.6 2.64 17.7 2.96 21.3 2.48 26.5 4.5 44
S. aureus 9.4 2.25 15.4 1.95 17.8 2.08 19.3 2.71 48
S. galifaeus 11.0 3.08 16.9 2.05 20.2 1.02 22.3 1.5 13
L. MARYUT
S. niloticus 11.1 2.52 22.1 1.78 26.5 2.32 30.7 13
S. nil0 ticuP 9.2 20.5 25.7 28.8
S. niloticusb 8.4 21.2 29.2 32.7
S. aureus 11.1 1.92 16.8 1.6 17.9 2.77 17
S. galilaeu@ 8.3 21.6 25.3 27.7
T. zilliP 7.2 13.3 18.4 21.1
L. QARUN
T. zilii’ 6.1 13.5 18.0 19.7

* Jensen (1957).
b El Zarka et al. (1970).
’ El Zarka (1961).

A comparison of our estimates of growth for S. niloticus and S. aureus shows

that, whereas the growth rate over the first year is very similar for both species,
beyond this the growth rate of S. aureus slows down appreciably. Applying
Students t-test showed that whilst there was no significant difference between
the sizes attained by S. niloticus and S. aureus after the first year in either L.
Manzalla or L. Maryut, there were significant differences between the two in
all subsequent years at both sites. As the size of the first maturity (Table IV)
appears to coincide with the size at the end of the first year this may indicate
that S. aureus channels proportionally more energy into reproduction than
S. niloticus. The final size of S. aureus is also probably smaller; the largest we
have recorded was 24 cm from Lake Manzalla compared to the largest S. nilo-
ticus of 32 cm, which approximately coincides with estimates of maximum
size given by the fisherman although they suggest S. niloticus can grow up to
40 cm.

TABLE IV

Minimum total length at first maturity for tilapia species in Lake Manzalla, Egypt

S. niloticus S. aureus S. galileus T. zillii

Male (cm) 17.4 11.3 14.8 11.7

Female (cm) 16.3 13.0 - 11.4
 343

In S. niloticus the mean size of the males was greater than that of the fe-
males by 0.6,0.4 and 1.7 cm for the first 3 years respectively but the dif-
ferences were found not to be significant. For S. aureus the females were on
average larger than the males for the first 2 years by up to 1 cm whilst in the
third year the reverse was true but none of the differences proved to be sig-
nificant.
The results of Jensen (1957) suggest that the growth rate of S. gulilaeus is
similar to that of S. niloticus whilst that of T. zillii is substantially lower. From
the present results the rate of S. galilaeus in L. Manzalla is slightly lower than
that of S. niloticus whilst still in advance of S. aureus. It is particularly notable
that the growth rates of T. zillii in L. Maryut (Jensen, 1957) and L. Qarun
(El Zarka, 1961) are so similar since the salinity at Maryut is around 5” lo,,
whilst that of Qarun is at least 20°/oo. There is no apparent inhibitory effect
of salinity on growth rate for this species over these ranges.
The estimates of mean length at the end of the first year included those
fish where the initial mark is a juvenile ring which can be taken as fish proba-
bly produced late in the spawning season and therefore of too small a size
to mature the following April, forming only the characteristic faint ring
instead of the more marked maturation ring. Such fish do not indicate the
true growth potential of the species over a full year which is of some interest
from the point of view of culture. For example, the mean size of the S. nilo-
ticus population after the first winter in L. Maryut was 11.1 cm (Table III)
but after excluding those with juvenile rings this mean becomes 12.9 cm and
can be interpreted as the mean length of fish spawned in the major part of
the spawning season which had grown to sufficient size to mature the follow-
ing year.

REPRODUCTION

Spawning

Spawning of tilapia in Egypt is seasonal owing to the distinct winter period.

In the delta area it may extend from April to November but with a peak in
early summer (El Saby, 1951). In Israel a similar pattern was found for S. gali-
laeus whilst the season for S. niloticus (although more probably S. aureus)
extended into May but may have subsided thereafter, although the findings
were not conclusive (Ben-Tuvia, 1959). The data in Table I suggest that S.
niloticus has a relatively restricted spawning season with a peak in April to
May whilst that of S. aureus is more generally protracted. Numbers in some
of the samples are rather small but there was abundant evidence of S. aureus
spawning in the shallow waters around the lakes in August although the
number of occupied nesting sites appeared to be fewer than in April. Nesting
behaviour was also observed in T. zillii in May and August and therefore this
species presumably also has a fairly lengthy spawning season.
From Table IV it appears that S. niloticus matures at a larger size than
344

S. aureus and the other two species. In addition, although it is difficult to

ascertain how frequently the fish spawn during the season, one female S.
aureus has been found with eggs in the mouth and mature ovaries in April
and it is not infrequent to find individuals of either sex with double rings at
a particular locus, both of these facts indicating that S. aureus can spawn at
least twice during the spawning season. There is no such evidence for S. nilo-
ticus, although more southerly populations from L. Nasser and the Nile at
Khartoum do give indications of more than one spawning per year (Latif,
and Rashid, 1972; Babiker and Ibrahim, 1979). All of these features point to
the fact that S. aureus expends rather more energy on reproduction than
S. niloticus, which may possibly explain the reduced growth rate beyond the
first year. There are also indications that S. aureus spawns in different areas
from S. niloticus which is an important point with respect to their specific
distinctiveness. S. aureus can be seen in shallow-water weedy areas with nests
of saucer shaped depressions of some 40 cm diameter and as little as 1 m
apart. S. niloticus was not seen in such locations and, according to the
fishermen, these spawn in deeper more open water locations, which indeed
reflects the generally preferred habitats of the two types.
The spawning cycle in lakes by no means reflects the cycle in a fish pond
since tilapia, at least in tropical regions, often mature at a smaller size and
spawn more frequently in ponds. However, observations at the Government
fish farm at El Manzalla, close to the shores of the lake, showed no sign of
reproductive activity in February and even in late April gonads were only in
the developing stage compared to lake fish which have commonly reached full
maturity by this time. Of a sample of fish from the farm in August only
S. aureus showed signs of reproductive activity but S. niloticus was under-
represented in the sample. The indications are, then, that the spawning season
in the ponds commences rather later than in the lake. This could be due to
greater fluctuations in temperature in the smaller water body of the ponds
and also, during the winter, to the generally lower pond temperatures com-
pared to those of the lakes.
Observations on the four tilapia species in Israel indicate that at 2O”C,
display and breeding behaviour begins to occur whilst at 22”C, spawning
itself can take place (Fishelson, 1966). In L. Manzalla the recorded monthly
maxima and minima are X.5-19.1 in March and 20.5-25.7”C in April
(Wahby et al., 1972) with similar values in L. Maryut (Aleem and Samaan,
1969) which coincides well with the observed commencement of the spawn-
ing season. For much of the early part of the year the temperature of the
Manzalla fish ponds is below the critical 22°C and in fact the monthly aver-
ages do not exceed 20°C until May (Eisawy et al., 1974). This goes some way
to explaining the delay in the start of the spawning season in the ponds
although an additional feature is that, in Egypt, all tilapia are stocked from
the wild each season, merely being allowed to colonise the ponds as they
enter with the water as the ponds are filled. The fish may therefore react as
wild individuals rather than cultured and not show the truncated life cycle.
 345

However, the relatively short breeding season must simplify the problem of
population control in Egyptian ponds compared to those of tropical Africa.

Fecundity

The mature ova in the ovaries of ripe female fish were counted and related
to body length for S. niloticus and S. aureus (Fig. 3). The eggs in each case
were orange-yellow in colour, rather elliptical in shape and some 2.5 mm
across at the longest point. Ripe ova can range from 1.94 to 2.95 mm (Latif
and Rashid, 1972). The data for both species followed the same pattern and
thus were pooled to give a relationship described by:
log F = log 2.14 + 2.25 log SL
or
log F = log 1.33 + 2.23 log TL
where F = number of mature eggs, SL = standard length (cm) and TL = total
length (cm). This is in reasonable agreement with the relationship of F = 2.895
SL’-“’ found in S. niloticus at Khartoum (Babiker and Ibrahim, 1979), and,
furthermore, comparison with values from farmed fish (Ruwet et al., 1976)
suggests that these also may be similar. The fecundity of S. aureus and S. nilo-

1600

", IlOO-
TLz1.33 SL"'gg5
r
lOOO-

a 5. niloticus
200 1
x s. oul-eus
loo-

10 11 12 13 14 15 16 17
Standard length (c-X”)

Fig. 3. The fecundity of Sarotherodon niloticus and S. aureus in relation to body length.
346

ticus is more closely related to the square of the length rather than the cube
as is most commonly found in fishes. A similar conclusion emerged for S.
leucostictus where it was considered that the fecundity was really associated
with the size of the oral brood pouch which in turn was related to the linear
dimensions rather than the weight of the fish (Welcome, 1967). However, an
alternative explanation is that whilst in most fishes fecundity increases in
proportion to weight, that is to the cube of the length, in tilapia it appears
to be more directly related to the square of the length that is to W”*66.In this
case fecundity proportionately declines with weight or alternatively the
smaller fish produce more eggs per g body weight. Since tilapia respond to
adverse conditions by reproducing at progressively smaller sizes at which they
are also relatively more fecund, by putting their biomass into smaller packets
they can produce more eggs to enhance the chance of survival in those condi-
tions.
A comparison with the data of other authors shows that the fecundity of
S. gclilaeus is similar to that found for the above species whilst that of T.
&ii is somewhat greater (Lowe-McConnell, 1955; Ben-Tuvia, 1959).

Hybridization

Under certain circumstances, principally when males of one species are

stocked with females of the other, S. niloticus and S. aureus will hybridize
often to give a skewed sex ratio which on occasion can give 100% males
(Trewavas, 1965, Fishelson, 1966). Serious attempts have been made in
Israel to make this cross the basis for the production of single sex fry for
monosex culture (Pruginin et al., 1975; Mires, 1977). Difficulties arose, how-
ever, due to the variable response of different strains and to inadvertant con-
tamination of maintained and natural pure stocks during commercial opera-
tions as a result of which the reliability of the procedure has been under-
mined (Pruginin et al., 1975). Eventually it was considered that the only way
to safeguard their genetic make-up was to do the crosses in indoor laborato-
ries where full control could be exerted (Mires, 1977).
In the Nile delta area several hundred fish of the two species have been
examined for their meristic features, particularly the dorsal fin ray numbers
and colour characteristics, notably the completely barred tail in S. niloticus
and the red caudal edge in S. aureus and it is virtually certain that hybridiza-
tion does not take place. Only two possible hybrids have been found and
these came from a pond on the Serow fish farm. The fish do seem to behave
as good species and appear to have rather different breeding grounds and
habitats as mentioned previously. The genetic distinctiveness of the two
species is further emphasized by the work of McAndrew and Majumdar
(1983) of the University of Stirling who have shown that muscle proteins
from live and frozen fish collected at Lake Manzalla have definite differences
when separated electrophoretically. These differences did not appear in fish,
nominally of these species, collected in Israel. It would seem, therefore, that
 347

Egypt has pure stocks of both S. aureus and S. niloticus which should be
regarded as a highly valuable resource if handled with care. Pruginin et al.
(1975) suggest that the maintenance of pure stocks would be consider-
ably easier if some genetic marker could be obtained for one of the species.
Two “golden”, presumably melanin deficient, individuals of S. aureus were
found around Lake Manzalla. If stocks could be bred from parents of this
type a convenient genetic marker may have been found.

SALINITY TOLERANCE

Salinity is of particular importance in the Nile delta owing to the wide

variation in this factor. The salinity records for a variety of waters in lower
Egypt are given in Table V.
The problem may be particularly acute where fish culture is attempted on
land where inadequate drainage rendered the soil unusable to agriculture
owing to accumulated salt which is a fairly common feature of the delta. The
canals and drains are the main waterways of the delta irrigation system and
also, potentially, the major water sources for fish farms. The drains tend to
be more saline than the canals (Table V). Shallow water fish ponds such as
are commonly used in Egypt often have salinities of 12-16°/oo, derived partly
from the soil and partly due to evaporation, even though their water sources
may be less than lo/ oo. Around Lake Manzalla shallow ponds may approach
the salinity of seawater, whilst even in the larger, established Manzalla Govern.
ment fish farm with deeper ponds the salinity is significant (Table V) and
may reach ll”/oo on occasions (Eisawy et al., 1974).
There is no doubt that T. zillii is the most salt tolerant of the tilapia species
in Egypt. El Zarka (1956) found that it survived well at 29°/oo and we have
seen it breeding in Lake Manzalla in water of 40°/oo NaCl. It is also the only
tilapia found along with grey mullet and gilthead bream (Sparus auratus) in
the saline ponds around this lake. Furthermore, it is the only tilapia surviving
in Lake Qarun, a lake which has been increasing progressively in salinity since

TABLE V

Conductivity and salt content of some major waters of lower Nile

Location Conductivity Total ions

(ClS x 103) (ppt NaCI)

R. Nile (Cairo) 0.39 < 0.1

L. Maryut 5.9-8.7 2.6-4.0
L. ManzaIla 4.3-40 2.0-40.0
ManzalIa fiih farm 7.7-11 3.0-7.0
Maryut canal 1.1-3.6 0.2-1.6
Maryut drain 4.1-10.1 1.8-5.7
L. Qarun 32 21.5
Sea 48 36
348

the 1920’s (Table V) and where at present the salinity can be up to 31°/oo in
the summer. It is notable that the growth rates of T. zillii in Lake Maryut and
Lake Qarun are not very different (Table III) despite the rather disparate sali-
nities. This emphasises the wide optimum tolerance range of T. zillii within
which the rate of growth is not depressed.
S. gulileus is generally reputed to have a high salinity tolerance although
precise information is scarce. The species has disappeared from Lake Qarun
whilst T. zillii persists, suggesting a lower tolerance than this species, a con-
clusion also suggested by the fish farmers of Lake Manzalla who indicated
that S. gulilueus disappeared from the areas of the Lake where the salinity
approaches that of seawater, whilst T. zillii is still found there. It has, however,
been shown that there is little difference in the growth of small 5’. galilueus
in tanks of freshwater, and 25 and 50% seawater (Chervinski, 1961a).
The relative salt tolerance of S. aureus and S. niloticus is difficult to assess
because of previous confusion in their identification. In tanks and earth ponds
S. uureus can grow equally as well in freshwater and water of 6.5,10.0 and
15.5°/oo NaCl although the mortality at the higher concentration ranged from
17.5-27.5% with a mean of 20.8% compared to an average of 8.3% in the
freshwater control (Chervinski, 1961b, 1966). It can spawn in 50% seawater
although larval survival is relatively low at this concentration. Moreover the
young survive direct transferral from fresh to 60% seawater much less well
than larger fish and, therefore, do not seem so well adapted to high salinity
as the adults (Chervinski, 1961b and c).
Unambiguous data on S. niloticus are very scarce but this species is probably
a true freshwater form since it persists along the length of the Nile whilst
S. uureus is confined to the delta. S. niloticus is another species which dis-
appeared from Lake Qarun probably when the salinity reached some 16°/oo
(S. El Zarka, personal communication, 1979). In Lake Manzalla, which has
connections to the sea, there is a salinity gradient from less than 1o/oo in the
south to full strength seawater in the north west, where it borders the sea.
Along this gradient S. niloticus is the first species to disappear and S. uureus
seems more persistent although less so than S. gulilueus and T. zillii (D.
Toews and T. Day, personal communication, 1979). The species is caught
around El Gamil where the salinity can reach 6°/oo and there is a thriving
population in Lake Maryut where the salinity is up to 5°/oo. Similarly S.
niloticus is found in the Manzalla fish farm at 7°/oo and mature S. niloticus
and S. uureus were stocked in ponds close to the Dead Sea where the salinity
varied from 3.7-14.9°/00 although their relative performance was not indi-
cated (Fishelson and Popper, 1968). In the last case, hybrids of the same
species also tolerated high salinities and seemed unaffected by the high con-
centrations of sulphates and manganese in the water. Therefore the effective
salinity tolerance for populations of S. niloticus might be between 5-10°/oo
whilst that for S. uureus could be higher, perhaps up to 15O/,,.
Fishelson (1980) found that, for experimental purposes, T. zillii, S.
mossumbicus, S. gulilueus and S. niloticus could all be acclimated to full
 349

strength seawater. However, S. niloticus proved least adaptable since they

showed highest mortalities in hypertonic water and could not become so well
adapted to hypersaline water as the other three species. Hybrids of S. nilotica
and S. aureus could become acclimated to these higher salinities and the in-
ference of this is that the salinity tolerance of S. aureus may be higher than
that of S. niloticus. Only 2’. zillii and S. mossambicus would spawn in seawater
and, in fact, T. zillii could be kept in up to 150% seawater. These observations
confirm the conclusions arrived at above on the relative salt tolerances
of the four species from the Nile Delta. However, the disappearance of S.
niloticus and S. galilaeus from Lake @run and the Great Bitter Lakes
(Bayoumi, 1969) at salinities well below those to which they can be acclimated
emphasises that individual tolerance limits do not define those for the main-
tenance of a population whilst the optimum limits for growth can be even
more restricted.

DISCUSSION AND CONCLUSIONS

It is apparent that S. niloticus grows at a faster rate than S. aureus in the

natural environments examined. This may possibly be because it commits less
energy to reproduction since it has a less protracted reproductive period and
may spawn more frequently. There may, however, be an intrinsic difference
between them. The salinity tolerance of S. niloticus appears lower than that
of S. aureus. These features indicate that at lower salinities S. niloticus may
give a better.response to culture than S. aureus whilst S. aureus may be pre-
ferable under slightly more saline conditions. It is, of course, difficult to
extrapolate from a lake to a pond environment but comparisons with fish in
the Manzalla fish farm have shown that spawning is seasonal, as in the lake,
and indeed may be a little later in commencing and that there was more
evidence of spawning activity in S. aureus than S. niloticus in August.
Of the other tilapia, the lower growth rate and higher fecundity of T. zillii
reduces its usefulness for culture. However, its high salinity tolerance does
mean that it can make some contribution to saline culture systems as it does
already in mixtures with the grey mullets, Mugil cephalus, Liza ramada and
gilthead bream, Sparus auratus in shallow ponds in the northern coastal
region where the water is close to full-strength seawater. In addition, of
course, it also fills a niche which would otherwise be unoccupied, by feeding
on higher plant materials.
S. galilueus grows well in the lakes and has a high salinity tolerance but
has generally been regarded as disappointing in culture (sarig, 1955; Lemas-
son, 1957).
The lack of hybridization between S. niloticus and S. aureus under natural
conditions in Egypt means that these should be regarded as a highly valuable
genetic resource providing pure parental stocks for the production of mono-
sex hybrids. They should be handled with great care to avoid the inter-breed-
ing, particularly between hybrids and parental stocks. This point is also true
350

for the rest of Africa and the precise identification of the stocks to be used
in any form of culture is crucial. The existing widespread transport of stocks
whose original provenance and genetic background are uncertain can lead to
breakdown of local species differences and will certainly make the job of the
selective fish breeder so much more difficult when the critical hurdle of true
domestication of tilapia is approached.

ACKNOWLEDGEMENTS

This work was carried out under the auspices of Atkins, Land and Water
management, Cambridge, Great Britain, in connection with the Nile Delta
Fish Farm Project for the Government of Egypt sponsored by the World
Bank and The Overseas Development Administration (Great Britain). Some
information was also collected during an aquaculture survey of Lake Manzalla
for James F. McClaren Co. (Canada) and UNDP.
We would like to thank Dr. Ali Katr for his help in planning the field
surveys and also Dr. E. Trewavas for her comments on the manuscript.

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