AQUATIC RESEARCH
E-ISSN 2618-6365
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Aspects of life history patterns of the cichlid fish Hemichromis
fasciatus Peters, 1857 from Okpara Stream, Northern Benin,
West Africa
Rachad Sidi Imorou , Alphonse Adite , Hamidou Arame , Pejanos Stanislas Sonon
Cite this article as:
Sidi Imorou, R., Adite, A., Arame, H., Sonon, P.S. (2020). Aspects of life history patterns of the cichlid fish Hemichromis fasciatus Peters, 1857 from
Okpara Stream, Northern Benin, West Africa. Aquatic Research, 3(2), 110-123. https://doi.org/10.3153/AR20010
University of Abomey-Calavi, Faculty of
Sciences and Technics, Department of
Zoology, Loboratory of Ecology and
Aquatic Ecosystems Management,
BP: 526, Cotonou, Benin
ORCID IDs of the author(s):
R.S.I. 0000-0001-6910-0059
A.A. 0000-0002-2255-4464
H.A. 0000-0002-0039-7787
P.S.S. 0000-0003-3810-7623
Submitted: 28.12.2019
Revision requested: 08.02.2020
Last revision received: 12.02.2020
Accepted: 18.02.2020
Published online: 22.03.2020
Correspondence:
Alphonse ADITE
E-mail: alphonseadite@gmail.com
©Copyright 2020 by ScientificWebJournals
Available online at
http://aquatres.scientificwebjournals.com
110
ABSTRACT
The piscivorous cichlid Hemichromis fasciatus dominated the Okpara Stream of Oueme River in
Benin, and numerically made 29.49% of the fish community. Some aspects of the reproductive
biology of this fresh/ brackish water cichlid were examined to evaluate the spawning patterns and
establishment of the species in Okpara Stream. Hemichromis fasciatus individuals were sampled
monthly during 18 months from December 2015 to May 2017 with seine, cast nets, gillnets, hooks
and traps. Among the 2,818 individuals sampled, 55.43% were females and 37.33% were males
with a sex-ratio of 1.49:1. The species showed low fecundities ranging between 113 and 1,716
oocytes. Length at first sexual maturity (L50) in Okpara Stream were reduced, 9.22 cm-TL (females) and 10.95 cm-TL (males), indicating an early maturation. The ovarian structure give evidence of multiple spawning and the species reproduces all seasons. In Okpara Stream, Hemichromis fasciatus exhibited a life history strategy between “r” and “K” selection, yet, close to “r”
selection. Because the length at first capture (Lc50 = 8.78 cm) is lower than sizes at first sexual
maturity, stocks of spawners could be reduced and leading to low recruitment.
Keywords: Life-history, Hemichromis fasciatus, Okpara Stream, Multiple spawning,
Recruitment, Early maturity
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Introduction
Hemichromis fasciatus (Actinopterygii: Perciformes: Cichlidae) is a widespread African piscivorous cichlid dwelling
most inland waters such as freshwater lakes, brackish waters,
estuaries, rivers, streams floodplains etc. because of its high
tolerance to environmental conditions (Leveque and Paugy,
2006, Kuela 2002, Paugy et al., 2003, Kantoussan 2007,
Montchowui et al., 2007 Sidi Imorou et al., 2019a). Like
most African cichlids and despite its reduced abundance, the
species showed great fisheries importance in the most water
bodies. As colourful species, Hemichromis fasciatus is utilized as ornamental fish and raised in the aquarium. The species has a laterally cylindrical compressed body mainly covered with cycloid scales ranging from 28 to 30 scales in lateral line (Lévêque and Paugy, 2006). Juveniles lack the orange-red coloring of adults and show traces of intervening
dark stripes. Subadult and adults in a state of stress also show
dark interstices. The maximum size observed is 204 mm
standard length (SL) (Lévêque and Paugy, 2006).
As feeding habit, H. fasciatus showed a moderate diet breadth
and foraged mainly on fishes, aquatic insects, zooplankton
and detritus. Unlike other cichlids, H. fasciatus is monogamous and protects eggs and fry but does not incubate orally
(Albaret, 1982). The eggs are laid on a block of rock, and are
fixed on a submerged support, in a clean place, sheltered from
the current and at a depth of 10 to 20 cm (Daget, 1956). Parental custody is highly developed in this species; both parents protect and accompany their offspring until they reach a
size of 2 to 3 cm.
In Benin, H. fasciatus is common and was recorded in almost
all water bodies and running waters where this piscivorous
cichlid showed reduced abundances varying between 0.37%
and 6% of the fish community, but constituted an important
fisheries resource (Jackson et al., 2013; Montchowui et al.,
2007). However, recent fisheries survey by Sidi Imorou et al.
(2019a) in the Okpara stream (North Benin), a tributary of the
Oueme River indicated that H. fasciatus dominated this
stream making numerically 29.49% of the fish assemblages.
Consequently, in the Okpara Stream, the species is intensively exploited and thus, displayed high commercial and
economic values in this northern running water.
Despite its high abundance and fisheries importance in the
Okpara stream associated with ongoing environmental
threats, nothing is known about the life history strategy and
the reproductive ecology of this piscivorous species in this
riverine water. This documentation is badly needed to characterize the species and to set up conservation, valorization
and a rational exploitation scheme that assure the ecosystem
balance.
The purpose of this fisheries study is to document some aspects of the reproductive biology of Hemichromis fasciatus
from Okpara Stream in order to provide basic knowledge
needed for management studies.
Material and Methods
Study Area and Sampling Sites
The study was performed in Okpara stream (200 Km), one of
the longest tributary of the Oueme River (510 Km). This river
sourced in the Northeast of Benin country at 450 m of altitude. Oueme River is the longest river and the more important
in term of fish species richness and fish production (Lalèyè et
al., 2004). Okpara stream is stretched between 8°14’9°45’ North and 2°35’-3°25’ East and belongs to the northern hydrographic network. The Benin northern region, is
characterized by a tropical climate with an alternation of dry
season (November - April) and wet season (May - August)
with a flood period from September to October. Annual ambient temperature averaged 26.6°C and lower temperatures
(18°C) were recorded in December-January. Annual mean
rainfall is about 1200 mm with a peak (2100 mm) recorded in
July or August (Kora, 2006). Soils are ferruginous and alluvial and covered by a wooded savanna of Parkia biglobosa,
Khaya senegalensis, Vitellaria paradoxa, marshy meadows,
bamboo and fallow bushes (Dossou-Yovo, 2009). Commercial fisheries take place in Okpara Stream that was exploited
by the surrounding populations. Also, this stream supplies the
surrounding populations with drinking water from a dam built
by SONEB, the Benin water company. Furthermore, Okpara
Stream provides water for irrigated agriculture.
For this study, H. fasciatus individuals were collected in five
(05) sampling sites (Figure 1). These sites were chosen according to localities, accessibility, fisheries importance and
levels of sites degradation. Site 1 is situated in Perere Township at Okpara up stream and Site 2 is localized in Parakou
Township at Gadela village (Okpara up stream), at about 2
km from SONEB dam. Site 3 is located at Kpassa village
where a dam was built to serve as a source of drinking water
for the populations of Tchaourou and Parakou Townships and
surrounding villages. Site 4 is situated around Okpara downstream at Yarimarou village (Tchaourou Township) where
the dam withdraws its water. Site 5 is also located around Okpara downstream at Sui village (Tchaourou Township). At
the five collecting sites, samplings were done in the “aquatic
vegetation habitat” at the edge of the stream and in the “open
water habitat” exempt of vegetation, but characterized by a
relatively high depth (Sidi Imorou et al., 2019a).
111
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Figure 1. Okpara stream and sampling sites. Site 1= Perere Township, Site 2 = Gadela village (Parakou Township), Site 3=
Kpassa village (Tchaourou Township), Site 4= Yarimarou village (Tchaourou Township), Site 5 = Sui village
(Tchaourou Township).
Okpara Stream displayed a relatively adequate water quality
with moderate temperatures ranging between 25 to 30.1°C,
depths between 17 to 1080 cm and low transparencies varying between 10 and 78.1 cm. Dissolved oxygen concentrations ranged between 0.44 and 5.66 mg/l and the percentages
of saturation were moderate to low and varied from 6.37 to
112
75.3%. The water was acid or alkaline with pHs ranging between 6.4 to 8.1 (Sidi Imorou et al., 2019a). As results, and
based on the physicochemical features recorded, the Okpara
stream is relatively suitable for a high primary production and
the prominence of the fish fauna.
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Fish Sampling
Hemichromis fasciatus samplings were done once a month
from December 2015 to May 2017 at all sites. Collections
were done in open water with experimental gill net and in
aquatic vegetation and with seines (Winemiller, 1992; Adite
et al., 2013; Gbaguidi and Adite, 2016, Sidi Imorou et al.,
2019a). Also, fish samplings were directly made in fishermen
artisanal captures with gill net (50 m x 1.30 m, 2.5 mm-mesh;
50 m x 1.30 m, 30 mm-mesh; 50 m x 1.30 m, 35 mm-mesh).
Fish species were identified using references such as Lévêque
and Paugy (2006). After collection, fishes were preserved in
10% formalin and then transported to the Laboratory of Ecology and Management of Aquatic Ecosystem (LEMEA)
where they were kept in 70% ethanol to make easier biological observations (Murphy and Willis, 1996).
Laboratory Procedure
In the laboratory, fishes were removed from the ethanol and
measured for total length (TL) and standard length (SL) to a
nearest 0.1mm with a digital caliper, and then weighed to the
nearest 0.1g with an electronic scale. Before dissection, sexes
were identified by examination of anal and genital opening
(Gbaguidi and Adite, 2016): males possess a tinny sexual
opening that also serves as anal orifice whereas females possess two (2) distinct orifices. After dissection, gonad weight,
gonad length and width were recorded, and stage of maturity
(I, II, III, IV, or V) in both males and females were evaluated
using the gonad maturation scale described by King (1995) :
(1) stage “immature” testis are simple translucent filaments
and ovaries are not developed, they are small and translucent.
Oocytes are not visible to the naked eye; (2) stage “Beginning
of maturation” During development, testicles are yellow or
ocher color and ovaries are opaque orange with opaque oocytes and visible to the naked eye; (3) “Advanced maturity”
testis are large and opaque white, ovaries are mature and fills
the ventral region of the abdominal cavity with translucent
eggs large and round; (4) stage “Mature” corresponding to
Ripe testis, fully developed and white milt was expelled by
gentle pressure on the abdomen. For females, Ovaries are mature and oocytes are ready to be expelled; (5) expelled sperm,
soft testes without milt, and expelled oocytes, narrowing of
the ovaries with some residual eggs.
The ovaries and testicles are then preserved in 90% alcohol.
Oocytes from mature ovaries were then separated using forceps and then enumerated. Oocytes diameters were recorded
using a binocular loupe. Batch fecundity was estimated as the
total number of oocytes in a ripe ovary. H. fasciatus oocytes
showed an ovoid form comprising with two dimensions (d1
and d2), measured with a calibrated eyepiece micrometer
Research Article
mounted to a dissecting stereomicroscope. Therefore, the theoretical diameters (d) utilized to construct the frequency histograms of egg diameters were the geometric means (d) of d1
and d2, expressed as following:
𝑑𝑑 = �(d1 × d2) (Adite et al., 2017)
Data Analysis
Data collected was recorded in Excel spreadsheet and analyzed with SPSS software version 21.0. The length of the
smallest mature specimen was the basis of the categorization
of the different life stages (Juveniles, subadults and adults)
and the length at first sexual maturation (TL50) for H. fasciatus (male and female) was estimated as the length at which
50% of the individual were mature as predicted by the formula of Ghorbel et al. (1996):
𝑷𝑷 =
𝟏𝟏
𝟏𝟏+𝒆𝒆−(𝒂𝒂+𝒃𝒃𝒃𝒃𝒃𝒃)
,
Where P is the pourcentage of individual mature, TL the total
length, a and b are constants. The length at first sexual matu−𝑎𝑎
ration were determined by: 𝑇𝑇𝑇𝑇50 =
𝑏𝑏
The sex ratio was computed as the ratio between the number
of males and the number of females. In order to evaluate the
spawning periodicity, the gonadosomatic indexes (GSI) were
calculated on the monthly basis for males and females following the formula:
GSI = (gonad weight / body weight)*100.
The batch fecundity was estimated for ripe ovaries, and the
relationship between fecundity (F) and Total length (TL) was
determinate following the curvilinear formula:
F = aTLb, where a and b are the parameters of the regression
In addition, the linear relationship between fecundity (F) and
body weight (W) was examined following linear regression:
F = bW + a, with a is the intercept, and b the slope.
For each season (flood, dry, wet), a mature gonad with a highest GSI was randomly chosen and frequency distributions of
oocyte diameters were plotted to examine the trends of egg
maturations and weather or not there was production of multiple cohorts per spawning season. The mean values of oocyte
diameters and gonadosommatic index (IGS) were given with
their standard deviation (mean ± standard deviation). Gender
percentages (sex ratio) and length at first sexual maturity
were compared using the χ2 test. Multiple comparisons of IGS
averages, and oocyte diameter were performed using a oneway ANOVA based on sex and seasons. The differences are
considered significant at p < 0.05. All analyze were done by
SPSS Software version 21.
113
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Results and Discussion
Population Structure and Sex Ratio
In Okpara Stream, a total of 2,818 individuals of Hemichromis fasciatus were sampled from December 2015 to May
2017. This abundance represented 29.49% of the fish assemblages making this piscivorous cichlid, the dominant species
of Okpara Stream fish community. In this stream, H. fasciatus
exhibited a bimodal size distributions dominated by individuals of total length ranging between 9 -11 cm (Figure 2).
Larger size (˃11 cm) abundances were reduced.
Of the 2,818 H. fasciatus specimen, 55.43% (1562 individuals) were females, 37.33% (1052 individuals) were males and
7.24% (204 individuals) were unidentified. This corresponded to a sex-ratio of 1:1.49 that is in favor of females.
This sex-ratio is significantly different to the theoretical sex
ratio, 1:1 (χ2= 99.503 p< 0.05) (Table 1). Also, the sex ratio
varied significantly with seasons with females dominating the
dry, wet and flood fish assemblages (Table 2).
Evolution of Gonadosomatic Index (GSI) and Spawning
Time
Figure 3 showed the variation of the gonadosomatic index of
H. fasciatus during the sampling period. Overall, the GSI
ranged from 0.001 to 15.202 and averaged 3.186 ±2.64 for
the females and there were significant difference (F18,1544=
2.588; p< 0.05) monthly variations. In males, the GSI values
were reduced and ranged between 0.01 and 1.70, with a mean
of 0.6 ±0.007. Like the females, the GSI of males showed
significant (F18,1052 = 8.144; p< 0.05) monthly variations.
Mature individuals (Stage 2, 3 and 4) are present all months
round with the highest proportions obtained in April
(78.88%), May (84.54%), June (82.29%), July (80.32%) in
males (Figure 4) and May (98.39%), June (91.56%), July
(77.52%) and August (75.00%) among females (Figure 5).
According to seasons, the wet and the flood were most favorable for the maturation of fishes with 96.52% and 95.82% respectively mature individuals.
Size at Sexual Maturity
Individuals of H. fasciatus with gonads at stages 2, 3 and 4 of
the maturation scales were considered as mature and used to
estimate the size at maturation (SSM). For the male, the
smallest mature individual measured 8 cm-TL and weighted
8.2 g, and the smallest mature individual for females measured 6.4 cm-TL and weighted 4.68 g. Percentage of ripe gonads reached 100% at sizes 14-17 cm for females and 15-17
114
Research Article
cm for males. Figures 6 and 7 illustrated the sigmoid curves
generated by the plot of ripe gonad percentages against total
length, respectively for males and females. The sigmoid
model showed that the sizes at sexual maturity for male and
female were 10.98 cm -TL and 9.22 cm-LT, respectively. The
sizes at maturity are not significantly different (χ²= 0.048475,
p> 0.05) between males and females. The regression’s equations were:
-
𝑃𝑃𝑓𝑓 =
𝑃𝑃𝑚𝑚 =
1
1+𝑒𝑒 −(−4.9345+0.5351𝑇𝑇𝑇𝑇)
1
1+𝑒𝑒 −(−6.1074+0.5558𝑇𝑇𝑇𝑇)
; r = 0.95 (for females)
; r = 0.91 (for males)
Ovarian Structure and Oocyte Sizes
Like in most cichlids, Hemichromis fasciatus possesses a pair
gonads (Moyle, 1988; Gbaguidi and Adite, 2016). Eggs
showed an ovoid shape and diameters varied from 0.800 mm
to 1.5297 mm with a mean of 1.10±0.11mm. Overall, the oocytes exhibited seasonal variations of diameters that ranged
between 0.8944 - 1.5297 (mean= 1.1280±0.1), 0.949 - 1.510
(mean= 1.192±0.12) and 0.80 - 1.342 (mean= 0.983±0.08)
during wet, flood and dry. Egg diameter frequency distributions by seasons showed unimodal distributions (Figure 8).
Absolute and Relative Fecundity
Of the 2,818 individuals of H. fasciatus sampled, 821 mature
females were recorded and were used to estimate batch fecundities. In this study, H. fasciatus showed batch fecundity
ranging between 113 oocytes for an individual of TL = 8.9
cm (W = 14.4g, GSI = 2.08) to 1716 oocytes for an individual
of TL = 9.7 cm (W = 19.5g, GSI = 5.64) with an average
value of 857 ± 284.78 oocytes for the fish assemblages. Relative fecundity varied from 15.21 oocytes/g (15 <TL <17) to
71.015 oocytes/g (7 <TL <9) (Table 2). Power curve fitted to
total length (TL) and fecundity (F) scatter plot showed that
batch fecundity significantly increase with TL (r = 0.93,
p<0.05) (Figure 8). Also linear regression fitted to body
weight (W) and batch fecundity scatter plot showed that fecundity significantly (r = 0.89, p<0.05) increase with body
weight (Figure 9). Regressions equations were as follows:
F = 16.369TL1.8637 (r = 0.93, N = 821) (Figure 9) and
F = 22.017W + 715.87 (r = 0.89, N = 821) (Figure 10)
With: F= Fecundity (Number of oocytes); TL= Total length
(cm); W= Body weight (g); N= Number of mature females;
r= correlation coefficient
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Table 1. Seasonal variations of the sex-ratio of H. fasciatus collected from December 2015 to May 2017 in Okpara Stream
(North Benin)
Seasons
Dry
Wet
Flood
Total
Males
590
422
40
1052
Females
693
766
103
1562
Sex ratio (M:F)
1:1.18
1:1.82
1:2.58
1:1.49
χ2
χ2 = 8.2689, p < 0.05
χ2 = 99.609, p < 0.05
χ2 = 27.755, p < 0.05
χ2 = 99.503, p < 0.05
Table 2. Mean, range and relative fecundities by length classes of H. fasciatus collected from Okpara Stream (North Benin)
TL class
(cm)
7-9
9-11
11-13
13-15
15-17
Total
N
141
666
58
14
2
881
Mean body
weigth (g)
11.42
16.60
30.18
49.34
69.33
17.36
Mean ovary
weigth (g)
0.58
0.68
0.72
0.86
1
0.67
Mean
fecundity
811
854
897
951
1055
857
Fecundity
range
168-1716
113-1555
397-1622
540-1484
945-1164
113-1716
Num of
eggs/ovary gram
1398
1256
1246
11.06
1055
1272
Relative
fecundity
71.015
51.45
29.72
19.27
15.21
49.08
Mean
GSI
5.05
4.14
2.46
1.78
1.24
4.13
Figure 2. Size structure (TL) of H. fasciatus (n = 2,818) collected in Okpara Stream (North-Benin) from December 2015 to
Mai 2017.
115
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Figure 3. Variation of gonadosomatic index of H. fasciatus from December 2015 to May 2017 in Okpara stream (North Benin)
Figure 4. Percentage of gonads maturation stages in H. fasciatus males by months. ST1 = Immature stage; ST2 = Stage of
beginning of maturation; ST3 = Advanced ripening stage; ST4 = Maturation stage and ST5 = Post-spawned
116
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Figure 5. Percentage of gonads maturation stages in H. fasciatus females by months. ST1 = Immature stage; ST2 = Stage of
beginning of maturation; ST3 = Advanced ripening stage; ST4 = Maturation stage and ST5 = Post-spawned
Figure 6. Lengths at first sexual maturity (L50) of H. fasciatus females (L50: 9.22cm, TL) from Okpara Stream, North Benin
117
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Figure 7. Lengths at first sexual maturity (L50) of H. fasciatus males (L50: 10.98 cm TL) from Okpara Stream, North Benin
Figure 8. Seasonal Egg diameter frequency distribution of H. fasciatus from December 2015 to May 2017in Okpara stream,
North Benin.
118
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Research Article
Figure 9. Relationship between Total length and fecundity of H. fasciatus in Okpara stream, North Benin
Figure 10. Relationship between Body weight and fecundity of H. fasciatus in Okpara stream, North Benin.
119
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Sex-ratio, Maturation and Spawning Time
As reported by many authors such Poulet (2004) in teleost
fishes, one sex predominates over the other. This general
trend agreed with that recorded for the population of H. fasciatus in Okpara Stream where the sex-ratio (1:1.49) recorded
was in favor of females with 1 male for 1.49 female. In Owa
Stream in Nigeria Olurin and Odeyemi (2010) rather reported
for H. fasciatus population, a sex ratio 1:0.36 in favor of
males. The morphological traits, the growth rates and the behavioral characteristics of males/females as selective may
play in favor of one sex. Also, in general, females are more
sedentary and more vulnerable to fishing gears because of parental care behavior such as egg incubation and offspring protection at the spawning grounds (Atse et al., 2009).
As reported by Sidi Imorou et al. (2019b), Okpara Stream
showed favorable growth conditions for H. fasciatus that exhibited an isometric growth pattern (slope b: 3.0024) and a
relatively high condition factor (K) reaching 8.02. Therefore,
spawning was effective, successful and the species was perfectly established. Indeed, the current study revealed the presence of different life stages small juvenile, juvenile, sub-adult
and adult in the population of H. fasciatus suggesting that the
reproduction and the recruitment of fishes were successful
(Gbaguidi and Adite, 2016). This result was also confirmed
by the presence of all maturation stages in the gonads. Abundance of oocytes of big diameters (d > 1.05 mm) during the
dry season (7.29%), the wet season (49.79%) and the flood
period (70.12%) combined with the relative abundance of
ripe gonads (stage 3 and 4) indicated that in Okpara Stream,
H. fasciatus breeds all seasons. This finding agreed with that
reported by Albaret (1982) in Ivory Coast and by KwarfoApegyah and Ofori-Danson (2010) in Batonga Reservoir in
Ghana where H. fasciatus spawned all year with a peak recorded in wet and flood seasons (Kwarfo-Apegyah and OforiDanson, 2010). As it is the case for many tropical fish species,
the increase of water level during wet and flood seasons was
the favorable condition that stimulates gonad maturation and
led to spawning and recruitment (Albaret 1982; Wootton
1998; Lalèyè et al., 1995; Dadebo et al., 2003; Rutaisire and
Booth 2005; Adité et al., 2006).
Length at First Sexual Maturation (TL50)
In Okpara Stream, males of H. fasciatus reached their size at
first maturity at a total length (L50 = 10.98 cm-TL) higher than
that of females (L50 = 9.22 cm -TL). This sexual difference in
L50, mostly tardive in males was reported for several fish species such Chrysichthys nigrodigitatus and Chrysichthys auratus (Lalèyè et al., 1995), Clarias ebriensis (Ezenwaji,
2002), Labeo horie (Dadebo et al., 2003), Labeo victorianus
(Rutaisire and Booth 2005); Heterotis niloticus (Adité et al.,
120
Research Article
2006), Clarias gariepinus, Clarias ebriensis, Schilbe intermedius, Schilbe mystus, Synodontis schall (Chikou et al.,
2011), Sarotherodon galileus (Gbaguidi and Adite 2016) etc.
Genetic factors including sexual dimorphism of growth and
habitat conditions such as physicochemical variations and
food resource availabilities could greatly affect the size at
first sexual maturity (Wootton, 1998; Koné et al. 2011; Tembeni et al. 2014).
In this study, the size at first sexual maturity (L50 = 9.22 cm TL) recorded for H. fasciatus females agreed with that reported by Olurin and Odeyemi (2010) in Owa Stream in Nigeria and that reported by Albaret (1982) in Ivory Coast.
Also, in Batonga Reservoir in Ghana, Kwarfo-Apegyah and
Ofori-Danson (2010) reported similar L50 for H. fasciatus. In
contrast, in Ogun River in Nigeria, Adebissi (1987) found a
higher L50=10.4 cm-TL for H. fasciatus female. Though genetic traits could cause this differential L50 through breeding
tactics and biological characteristics, habitat ecological status, fishery condition, population structure and environmental conditions and stockascity could together influence the
length at first sexual maturity (Rutaisire and Booth, 2005). In
Okpara stream, the length at first capture (Lc50) for H. fasciatus is Lc50=8.78cm (Sidi Imorou et al., 2019c). This length
is lower than the length at first maturity (L50=10.98 cm for
males and L50=9.22 cm for females). This result suggested
that in Okpara stream, specimen of H. fasciatus were caught
before breeding for the first time. This situation could lead to
a depletion of the stock of this species in the river.
Power curve fitted to total length (TL) and fecundity (F) scatter plot showed that batch fecundity significantly increase
with TL (r= 0.93, p<0.05) (Figure 8). Also linear regression
fitted to body weight (W) and batch fecundity scatter plot
showed that fecundity significantly (r = 0.89, p<0.05) increase with body weight (Figure 9).
Fecundity and Breeding Strategy
In the current fishery survey, H. fasciatus showed batch fecundities ranging between 113 and 1716 oocytes (mean = 857
oocytes) with a relative fecundity of 49.08 oocytes / g of body
weight in Ivory Coast, Albaret (1982) recorded a highest fecundity that reached 2509 oocyte along with a reduced relative fecundity of 30 oocytes / g of body weight. In contrast,
in New Bussa kigera reservoir in Nigeria, Issa et al. (2005)
reported a lowest fecundity of 520 oocytes. Probably, these
spatial differences in fecundities were the results of differential environmental variabilities and differential availability in
food resources (Fagade et al., 1984; Issa et al., 2005). In this
study, batch fecundity exhibited significant (r= 0.93, p< 0.05)
curvilinear relationship with total length and was positively
correlated with body weight. Legendre and Ecoutin (1989)
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
and Atse et al. (2009) reported similar trends in some other
cichlids such as Sarotherodon melanotheron, Tilapia guineensis and Tylochromis jentinki from Ebrie Lagoon in Ivory
Coast.
With regards to ovarian structure, H. fasciatus showed moderate oocyte diameter averaging 1.10 ± 0.11 mm and varying
between 0.800 mm and 1.5297 mm. These diameter values
were lower than those reported by Albaret (1982) in the water
bodies of Ivory Coast and by Olurin and Odeyemi (2010) in
Owa Stream in Nigeria where the oocyte diameters reached
1.65 mm and 1.94 mm, respectively. Also, the gonads comprised oocytes at all stages of maturity regardless of month
(Figures 3 and 4), suggesting that H. fasciatus displayed multiple reproduction in the Okpara River with several offspring
cohorts during the same spawning season. These breeding
pattern is typical to most teleost fishes and cichlids in particular (Paugy et al., 2006).
In this study, H. fasciatus exhibited a relative low sizes at
sexual maturity 10.98 cm-TL (males) and 9.22 cm-LT (females), a low batch fecundity (F = 857 ± 284.78 oocytes), a
relative low eggs diameters (d=1.10±0.11mm) and reproduce
early. This piscivorious fish built nests in which eggs are laid
and had multiple layers and practices parental care for the
survival of its offspring (Stiassny et al., 2007). These biological characteristics indicate that H. fasciatus has a demographic strategy between selections "r" and "K" but much
closer to selection "r". This strategy optimizes the survival of
eggs, larvae and juveniles through the practice of parental
care with moderately early maturation associated with several
spawnings throughout the year (Winemiller and Rose, 1992).
Conclusion
The current study on the reproductive biology of H. fasciatus
gives valuable information on the spawning time, fecundity
and spawning strategy of this fish species in Okpara stream.
The favorable environmental conditions of Okpara stream
coupled with the all seasons breeding and the spawning strategy characterized by the high offspring survivorship (parental
care), the high accounted for the prominence in this ecosystem. However, values of size at first sexual maturity for both
sexes superior to the length at first capture require special attention in the sustainable management plan for this ecosystem.
Compliance with Ethical Standard
Conflict of interests: The authors declare that for this article they have no
actual, potential or perceived conflict of interests.
Research Article
Funding: This research was funded by the “Laboratory of Ecology and
Management of Aquatic Ecosystems” and by the authors.
Acknowledgments: We are grateful to the “Laboratoire d’Ecologie et de
Management des Ecosystèmes Aquatiques, Département de Zoologie, Faculté des Sciences et techniques, Université d’Abomey-Calavi” for providing us logistic assistances. We are also grateful to Okpara’s fishermen for
their help during the investigation. We express our gratitude to Amoussouga Illary and Mitobaba Aurel for their assistance in laboratory works.
References
Adite, A., Winemiller, K.O., Fiogbe, E.D. (2006). Population structure and reproduction of the African bonytongue
Heterotis niloticus in the Soˆ River-floodplain system (West
Africa): implications for management. Ecology of Freshwater Fishes, 15, 30-39.
https://doi.org/10.1111/j.1600-0633.2005.00119.x
Albaret, J.J. (1982). Reproduction et fécondité des poissons
d'eau douce de Côte-d'Ivoire. Revue Hydrobiologique Tropicale, 15, 347-371.
Adite, A., Gbaguidi, H.M.G.A., Ategbo, J-M. (2017). Reproductive biology and life history patterns of the Claroteid,
Chrysichthys nigrodigitatus (Lacépède: 1803) from a Manmade lake in Southern Benin. Journal of Fisheries and
Aquatic Science, 12, 106-116.
https://doi.org/10.3923/jfas.2017.106.116
Chikou, A., Laleye, P.A., Bonou, C.A., Vandewalle, P.,
Philippart, J-C. (2011). Tailles de première maturité et de
capture de six espèces de poisson-chat dans le delta de
l'Ouémé au Bénin (Afrique de l'Ouest). International Journal
of Biological and Chemicale Science, 5(4), 1527-1537.
https://doi.org/10.4314/ijbcs.v5i4.18
Atse, B.C., Konan, K.J., Kouassi, N.J. (2009). Biologie de
la reproduction du Cichlidae Tylochromis jentinki dans la lagune Ebrié (Cote d'Ivoire). Cymbium, 33(1), 11-19.
Dadebo, E., Ahlgren, G., Ahlgren, I. (2003). Aspect of reproductive biology of Labeo horie Heckel (Pisces: Cyprinidae) in Lake Chamo. Ethiopia. African Journal of Ecology,
41, 31-38.
https://doi.org/10.1046/j.1365-2028.2003.00404.x
Ethics committee approval: This study was conducted in accordance with
ethics committee procedures of animal experiments.
121
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Daget, J. (1956). La pêche à Diafarabé. Études
monographiques. Bulletin de l'Institut fondamentale de l'Afrique noire, série B, 18(1-2), 97.
Dossou-Yovo, E. (2009). Modélisation du fonctionnement
hydrologique dans le bassin versant de l'Okpara à l'exutoire
de Kaboua dans un contexte de changement global: contribution à la gestion intégrée des ressources en eau. Thèse d'Ingénieur Agronome, Faculté des Sciences Agronomiques,
Abomey-Calavi, Bénin. 106p.
Kantoussan, J. (2007). Impact de la pression de pêche sur
l'organisation des peuplements de poissons: application aux
retenues artificielles de Selingue et de Manantali, Mali, Afrique de l'Ouest. Thèse de doctorat, Agrocampus, Rennes,
195p.
Koné, N., Berte, S., Kraidy, A.L.B., Kouamelan, E.P.,
Koné, T. (2011). Biologie de la reproduction du Clupeidae
Pellonula leonensis Boulenger, 1916 dans le lac de barrage
de Kossou (Côte d'Ivoire). Journal of Applied Biosciences,
41, 2797-2807.
Ezenwaji H.M.G. (2002). The biology of Clarias ebriensis
Pellegrin, 1920 (Osteichthyes: Clariidae) in an Africa rainforest river basin. Fisheries Research, 54, 235-252.
https://doi.org/10.1016/S0165-7836(00)00291-5
Kora, O. (2006). Monographie de la commune de Parakou.
Cabinet « Afrique Conseil », 44p.
Fagade, S.O., Adebisi, A.A., Atanda, A.N. (1984). The
breeding cycle of Sarotherodon galilaeus in the IITA Lake,
Ibadan, Nigeria. Archives Hydrobiologia, 100(4), 493 - 500.
Kuela, J.M.D. (2002). Étude des peuplements ichtyologiques de la Comoé et des modes d'exploitation piscicole
dans la zone agro-sylvo-pastorale du projet GEPRENAF.
Mémoire de fin d'étude, Université polytechnique de Bobo
Dioulasso, 73p.
Gbaguidi, H.M.G.A., Adite, A. (2016). Reproductive ecology and establishment of naturally colonized tilapine Cichlid, Sarotherodon galilaeus (Pisces: Actinopterigii: Perciformes) from a man-made lake, Southern Benin, West Africa: implications for sustainable fisheries and aquacultural
valorization. International Journal of Fisheries and Aquatic
Studies, 4(3), 278-287.
Kwarfo-Apegyah, K., Ofori-Danson, P.K. (2010). Spawning and recruitment patterns of major fish species in Bontanga reservoir, Ghana, West Africa. Lakes & Reservoirs:
Research and Management, 15, 3-14.
https://doi.org/10.1111/j.1440-1770.2010.00418.x
Ghorbel, M., Jarboui, O., Bradai, M.N., Bouain, A.
(1996). Détermination de la taille de première maturité sexuelle par une fonction logistique chez Limanda limanda, Pagellus erythrinus et Scorpaena porcus. Bulletin INSTM, 3, 2427.
Issa, E.A., Olufeagba, S.O., Raji, A. (2005). Effects of
some environmental factors on the fecundity of Tilapia species (Family: Cichlidae) in Kigera Reservoir, New Bussa.
19th Annual Conference of the Fisheries Society of Nigeria
(FISON), 29th Nov. - 03 Dec. 2004, Ilorin, Nigeria.
Jackson, A.T., Adite, A., Roach, K.A., Winemiller, K.O.
(2013). Fish assemblages of an African river floodplain: a
test of alternative models of community structure. Ecology of
Freshwater Fish, 22, 295-306.
https://doi.org/10.1111/eff.12026
122
Research Article
Lalèyè, P., Chikou, A., Philippart, J-C., Teugels, G.,
Vandewalle, P. (2004). Étude de la diversité ichtyologique
du bassin du fleuve Ouémé au Bénin (Afrique de l'Ouest).
Cybium, 28(4), 329-339.
Lalèyè, P., Phillipart, J-C., Poncin, P. (1995). Biologie de
la reproduction de deux espèces de Chrysichthys (Siluriformes, Bagridae) du lac Nokoué et de la lagune de Porto
Novo au Bénin. Journal of African Zoology, 109(3), 213224.
Legendre, M., Ecoutin, J.M. (1989). Suitability of brackish
water tilapia species from the Ivory Coast for lagoon aquaculture: Reproduction. Aquatic Living Resources, 2, 71-79.
https://doi.org/10.1051/alr:1989009
Lévêque, C., Paugy, D. (2006). Les poissons des eaux continentales africaines: Diversité, écologie, utilisation par
l'homme. IRD Editions. 573p.
Aquat Res 3(2), 110-123 (2020) • https://doi.org/10.3153/AR20010
Stiassny, M.L.J., Teugels, G.G., Hopkins C.D. (éd.)
(2007). Poissons d'eaux douces et saumâtres de basse
Guinée, ouest de l'Afrique centrale, Collection Faune et Flore
tropicales. IRD, Publications scientifiques du Muséum,
MRAC, ISSN: 1286-4994
Montchowui, E., Niyonkuru, C., Ahouansou Montcho, S.,
Chikou, A., Lalèyè, P. (2007). L'ichtyofaune de la rivière
Hlan au Bénin (Afrique de l'Ouest). Cybium, 31(2), 163-166.
https://doi.org/10.4314/ijbcs.v2i2.39733
Murphy, B.R., Willis, D.W. (1996). Fisheries Techniques.
Second edition. American Fisheries Society, Bethesa, Maryland. ISBN-13: 978-1888569001
Olurin, K.B., Odeyemi, O.I. (2010). The reproductive biology of the fishes of Owa Stream, South-West Nigeria. Research Journal of Fisheries and Hydrobiology, 5(2), 81-84.
Paugy, D., Lévêque, C., Teugels, G.G. (2003). Faune des
poissons d'eaux douces et saumatres de l'Afrique de l'Ouest.
Tome 1. IRD Editions, MNHN, Paris, France & MRAC, Tervuren, Belgique, 457p.
Paugy, D., Lévêque, C., Duponchelle, F. (2006). La reproduction in Lévêque C., Paugy D. (2006). Les poissons des
eaux continentales africaines : Diversité, écologie, utilisation
par l'homme; IRD Editions, Paris, France, ISBN 2e édition :
2-7099-1589-8
Poulet, N. (2004). Le sandre (Sander lucioperca L.) : Biologie, comportement et dynamique des populations en Carmargue (Bouches du Rhône, France). Thèse de Doctorat,
Université de Toulouse III, France, 300 p.
Rutaisire, J., Booth, A.J. (2005). Reproductive biology of
ningu, Labeo victorianus (Pisces: Cyprinidae), in the Kagera
and Sio Rivers, Uganda. Environmental Biology of Fish, 73,
153-162.
https://doi.org/10.1007/s10641-004-5564-8
Research Article
Sidi Imorou, R., Adite, A., Arame, H., Chikou, A.,
Adjibade, N.K., Sonon, P.S. (2019a). Ichthyofauna of Okpara Stream, a Tributary of Oueme River, Benin, West-Africa. International Journal of Sciences, 8(5), 53-66.
https://doi.org/10.18483/ijSci.2052
Sidi Imorou, R., Adite, A., Sossoukpe, E., Abou, Y.
(2019b). Length-weight models and condition factors of
fishes from Okpara Stream, Oueme River, Northern-Benin.
International Journal of Forest, Animal and Fisheries Research, 3(3), 65-80.
https://doi.org/10.22161/ijfaf.3.3.1
Sidi Imorou, R., Adite, A., Sossoukpe, E., Adjibade, N.K.,
Arame, H., Sonon, SP. (2019c). Demographic parameters
and exploitation rate of five key fishes of Okpara Stream,
Oueme River, Benin, West Africa. International Journal of
Aquatic Biology, 7(6), 332-341.
https://doi.org/10.22034/ijab.v7i6.642
Tembeni, J.M., Micha, J.C., Mbomba, B.N.S., Vandewalle, P., Mbadu, V.Z. (2014). Biologie de la reproduction
d'un poisson chat Africain Euchilichthys guentheri
(Schilthuis, 1891) (Mochokidae, Siluriformes) au Pool
Malebo, Fleuve Congo (République Démocratique du
Congo). Tropicultura, 32(3), 129-137.
Winemiller, K.O., Rose, K.A. (1992). Patterns of life-history diversification in North American fishes: implications
for population regulation. Canadian Journal of Fisheries and
Aquatic Science, 49, 2196-2218.
https://doi.org/10.1139/f92-242
Winemiller, K.O. (1992). Fish assemblages across a complex, tropical freshwater/marine ecotone. Environmental Biology of Fishes, 3, 29-50.
https://doi.org/10.1007/BF00004783
Wootton, R.J. (1998). Ecology of Teleost Fishes. Kluwer
Academic Publishers [Fish and Fisheries Series, no. 24],
Dordrecht, The Netherlands. ISBN: 978-0-412-84590-1
123