Aquaculture and Fisheries 6 (2021) 424–431

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Aquaculture and Fisheries

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Reproductive cycle, sexual maturity and fecundity of Nemipterus furcosus

(Valenciennes, 1830)
Mustafizur M. Rahman a, b, *, Amira F. Samat c
a
Institute of Oceanography and Maritime Studies, International Islamic University Malaysia (IIUM), Kg. Cherok Paloh, Kuantan, Pahang, 26160, Malaysia
b
Department of Marine Science, IIUM, Jalan Sultan Ahmad Shah, Bandar Indera Mahkota, Kuantan, Pahang, 25200, Malaysia
c
Universiti Malaysia Perlis, Sungai Chuchuh, Padang Besar, Perlis, 02100, Malaysia

A R T I C L E I N F O A B S T R A C T

Keywords: Understanding the reproductive biology particularly sexual maturity, reproductive cycle and fecundity of fish is
Gonadal histology useful for fisheries management. Unfortunately, there is no information about reproductive biology of Nemipterus
Sexual maturity furcosus. This study addresses this deficit for N. furcosus. We conducted a 12-month study to understand the
Nemipterus furcosus
reproductive biology particularly the sex ratio, sexual maturity, fecundity and reproductive cycle of N. furcosus at
Oocytes diameter
South China Sea
the east coast of peninsular Malaysia. Results indicated that ratios of male and female were statistically similar up
to 19.0 cm TL (total length). Male outnumbered female above 19.0 cm TL (P < 0.05). The fecundity ranged
between 54,970 and 236,938 with a mean fecundity of 102,477 ± 43,580 (standard deviation). Male and female
fish reached first sexual maturation at almost same size, 15.8 and 15.6 cm, respectively. This information is
needed to determine the lowest acceptable harvest length of this fish under the management aim to enter all
female fish for reproduction minimum one time in their life. We estimated the reproductive period of N. furcosus
applying oocytes diameter, histological examination of ovaries and monthly mean GSI. All of these methods gave
a consistent result about spawning season, which appeared to be from March to July with peaks in March and
June. Based on this result, N. furcosus spawners can be protected during March to July for entering the repro­
ductive stock. Apart from artificial breeding, the results of this study might facilitate future efforts at fine-tuning
fishing effort for a proper management of N. furcosus.

1. Introduction
The genus Nemipterus encompasses more than 50 nominal species
and are distributed all over the Indo-West Pacific region (Russell, 1993).
Most species of this genus are normally found near the sandy or muddy
bottoms at depths from 5 to 80 m (Kerdgari, Valinassab, Jamili, Fatemi,
& Kaymaram, 2009; Russell, 1993). Generally, nemipterids are
commercially caught by bottom trawl. They are one of the most
commercially important groups of marine fishes particularly in the east
China sea (Tonnie et al., 2018). Due to their importance in fisheries,
many studies have been conducted regarding various aspects of biology
of nemipterids, particularly N. peronii (Said, Mohsin, & Ambak, 1994;
Wu, Weng, Liu, & Su, 2008), N. virgatus (Takahashi, Kobayashi, &
Suzuki, 1989), N. Bathibius (Granada, Masuda, & Matsuoka, 2004;
Takahashi et al., 1989), N. japonicus (Raje, 2002, Rajkumar, Rao, &
Kingsly, 2003; Manojkumar, 2004; Kerdgari et al., 2009; ElHaweet,
2013; Nettely et al., 2016) and N. randalli (Erguden, Turan, Gurlek,
Yaglioglu, & Gungor, 2010; Innal et al., 2015). Unfortunately, infor­
mation regarding reproductive biology of N. furcosus is very limited
(Amira et al., 2016; FAO, 2012, p. 209). Persada, Utami, and Rosalina
(2016) conducted a three-month (March–May 2015) study on a few
reproductive parameters (sex ratio, gonadosomatic index, facundity,
etc.) of N. furcosus landed at the Sungailiat archipelago fishery Port,
Indonesia.
Nemipterus furcosus, frequently called forktail threadfin bream, is one
of the most popular and economically important food fish in many Asian
countries, where it is an important coastal demersal fisheries resource
(Amira et al., 2016). It is distributed in numerous Asian and Australian
coasts. Due to high demand and market price, it is one of the chosen
fishes for Malaysian fisher and therefore, total production of N. furcosus
on the east coast of Malaysian peninsular is decreasing day by day
(Amira et al., 2016; FAO, 2012, p. 209). Presently, it is also a target fish
for aquaculture to meet the consumer demand. However, artificial
breeding of this fish is not successful yet due to insufficient knowledge

* Corresponding author. Institute of Oceanography and Maritime Studies, IIUM, Kg. Cherok Paloh, 26160, Kuantan, Pahang, Malaysia.
E-mail address: mustafizu.rahman@yahoo.com (M.M. Rahman).

https://doi.org/10.1016/j.aaf.2020.07.006
Received 2 March 2020; Received in revised form 3 July 2020; Accepted 8 July 2020
Available online 10 August 2020
2468-550X/© 2020 Shanghai Ocean University. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd. This is an open access article
under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
M.M. Rahman and A.F. Samat
regarding reproductive biology especially sexual maturity, fecundity,
gonadal change and spawning period.
Size at first sexual maturity (SFSM) is applied to estimate the lowest
acceptable harvest length of fish to enter female fish for reproduction
minimum one time in their life (Rahman, 2017). Understanding fecun­
dity is useful to evaluate the commercial possibilities of fish populations
(Tracey, Lyle, & Haddon, 2007). It helps estimating the revival potential
of overexploited fish populations (Hughes & Stewart, 2006). Rules and
legislations need to be applied to prohibit catching spawners during the
reproductive period for maintaining sustainability of the fish produc­
tion. Therefore, besides artificial breeding, understanding the repro­
ductive parameters of fish is crucial for fishery managers, who usually
depend on the SFSM, and the starting and duration of reproduction for
fisheries management (Jakobsen, Fogarty, Megrey, & Moksness, 2009;
Trindade-Santos & Freire, 2015). The lack of this information in some
countries like Malaysia often leads to overfishing and management
failure (Alves & Minte-Vera, 2012; Kjesbu, 2016). Unfortunately, in­
formation about the reproductive parameters of N. Furcosus is scarce,
particularly in Asian countries including Malaysia. This study sought to
contribute towards addressing the deficit of information by investigating
the sex ratio, reproductive cycle, sexual maturity, fecundity and the
correlation of fecundity with length and weight of N. furcosus for a
period of 12 months at the east coast of peninsular Malaysia. The in­
formation of this paper can be served as an important reference for the
N. furcosus management decision making.
2. Materials and methods
2.1. Sampling and measurement
All fish were sampled randomly from fishing vessels, which caught
fish at the east coast of peninsular Malaysia (Fig. 1) using trawl nets. We
used random sampling method for avoiding biasness in the homogenous
group of fish. After arriving the fishing vessels at the landing center, all
fish are kept in a pile on the clean concrete made floor for manually
sorting them based on size. We sampled a portion (approximately 10–20
kg) of the fish before sorting them out. Fish sampling was conducted
monthly over 12 months (Aug 2012 to Jul 2013) to incorporate a full
seasonal cycle. A total of 1446 forktail threadfin bream specimens were
collected for this research. Sample size in October and November were
comparatively low due to very rough coastal weather. After collection,
Fig. 1. Map showing N. furcosus capture location at the East coast of Peninsular Malaysia.
425
Aquaculture and Fisheries 6 (2021) 424–431
specimens were immediately iced and transported to the Marine Science
Laboratory, IIUM. Confirmation of species was done using an identifi­
cation key (Mohsin & Ambak, 1983). Total length (TL) and total body
weight (TW) were determined using a centimeter scale and a weighing
scale (precision 0.01 g), respectively. Gonad of each fish were collected
by dissection and were weighed using the same digital balance. Sex was
determined via the observation of gonads, by naked eye or by micro­
scope. Sexual dimorphism was checked by naked eyes. In each month,
randomly selected 50% (n = 25 (Aug), 20 (Sep), (9) Oct, 4 (Nov), 20
(Dec), 17 (Jan), 23 (Feb), 24 (Mar), 23 (Apr), 43 (May), 45 (Jun), 35
(Jul) of the total ovary were preserved in 10% phosphate buffered
formalin for histological examination and other ovaries (n = 24 (Aug),
19 (Sep), 9 (Oct), 4 (Nov), 20 (Dec), 16 (Jan), 22 (Feb), 23 (Mar), 23
(Apr), 42 (May), 45 (Jun), 34 (Jul)) were preserved in Gilson fluid to
estimate oocyte diameter. Gonadosomatic index (GSI), SFSM, fecundity,
size of oocyte and histological examination of ovary were estimated as
female reproductive parameters, while GSI and SFSM were estimated as
male reproductive parameters.
2.2. Estimation of gonadosomatic index, size at first sexual maturity,
fecundity and oocyte diameter
GSI was determined applying the formula, GSI (%) = GW BW × 100
with GW = gonad weight (g), BW = body weight (g). The SFSM was
estimated by plotting total length (X axis) versus gonadosomatic index
(Y axis) (Ohtomi, Tashiro, Atsuchi, & Kohno, 2003). It was further
calculated using the empirical equation, LogLm = − 0.1189 + 0.9157 ×
logLmax (Binohlan & Froese, 2009) with Lm = the SFSM, Lmax = the
maximum total length.
All ovaries preserved in Gilson fluid were used to estimate oocyte
diameter. Preserved (in Gilson fluid) ovaries with a GSI ≥ 2.2% were
considered as matured ovaries and were used for fecundity estimation. A
total of 104 ovaries were used for fecundity estimation. For this, a sub-
sample of 0.05 g was taken from each of 6 portions (the anterior, middle,
and posterior of both lobes) and counted number of eggs under a stereo
microscope. The fecundity (F) was calculated using the equation, F =
((Nss × Wo ) /Wss )with Nss = number of oocytes in 6 subsamples, Wo =
total weight the ovary, Wss = total weight of 6 subsamples.
The relationships of fecundity with body length (L) and fecundity (F)
with body weight (W) were determined using the equations, F = aLb and
F = aWb, respectively. The constants b and R2 were determined using a
M.M. Rahman and A.F. Samat Aquaculture and Fisheries 6 (2021) 424–431

linear regression on the logarithmic transformed equations LogF = log a opposite result was observed in May and June. Overall male N. furcosus
+ b logL and LogF = log a + b logW, respectively. ‘ (n = 877) was significantly higher (P < 0.05) in number compared to the
The size of oocytes was determined using a microscope connected female N. furcosus (n = 569).
with a digital camera mounted on a computer and software.

2.3. Histological examination of ovary 3.2. Fecundity

Four major steps of tissue processing (dehydration, clearing, infil­
tration, and impregnation and embedding) were done to process ovaries
according to Morrison (1990). Firstly, the ovaries were dehydrated by
running them with a series of alcohol in increasing concentrations from
50% to 100% and cleared in an automated tissue processor. The cleared
tissues were pre-embedded with toluene-paraffin and fully embedded in
paraffin wax. The embedded tissues were poured into a mold and cooled
on the cold plate. After cooling, they were sectioned at 6 μm thickness by
a microtome. The sliced tissues were put on slides and stained with
haematoxylin and eosin. The stained tissues were mounted in DPX
mountant and observed under a microscope connected with a digital
camera. Gonad maturation was categorized (six stages) by histological
inspection according to Bancroft and Stevens (1996).
2.4. Data analyses
The significant difference between male and female ratio was
determined through the χ2 test. A one-way ANOVA (significant at P ≤
0.05) performed to find the temporal variation of the GSI. ANOVA was
followed by Tukey test if the effect was significant. All data were
examined for suitability of parametric test before performing the
ANOVA.
The fecundity varied from 54,970 to 236,938 with an overall mean of
102,477 ± 43,580 (standard deviation). Although not reported here,
monthly mean fecundity of N. furcosus was statistically similar (P >
0.05). Fecundity (F) of fish increased with total length (TL) and total
body weight (W): F = 80.1 × TL2.374 (R2 = 0.94, P < 0.01, n = 104) and
F = 11.2 × 102 × W0.953 (R2 = 0.95, P < 0.01, n = 104) (Fig. 2).
3.3. Size of oocytes
The size of oocyte was highly variable and ranged from 0.09 to 1.18
mm in diameter. Fig. 3 represents the variation in the oocyte size over 12
months to incorporate a full seasonal cycle. Large-sized oocytes >0.5
mm in diameter were recorded mostly during January to August, with
highest sized group 1.11–1.20 mm during March to July. Small-sized
group <0.5 mm was recorded mostly during August to December. The
size of oocyte was related with the gonadosomatic index. The size of
oocyte from ovaries with an average gonadosomatic index of >2.12 was
almost always >0.5 mm in diameter. The size of oocyte from ovaries
with an average gonadosomatic index of <1.1 was almost always <0.5
mm in diameter. Once oocytes became ≥0.5 mm in diameter were
considered matured (Wu et al., 2008).

3. Results

3.1. Sex ratio

The sex ratio of N. furcosus was statistically similar up to 19.0 cm

total length (P > 0.05) (Table 1). Male significantly more than female
above 19.0 cm TL (P < 0.05). The ratio of male and female were sta­
tistically different in all months except April and July (Table 1). The
ratio of male was higher than female in January, February, March,
August, September, October, November and December, whereas the

Table 1
Ratio of female N. furcosus by size and month during August 2012–July 2013.
Male Female Ratio (proportion of χ2 value P value
female)

Size class (cm)

9.0–11.0 22 27 0.55 0.510 0.475
11.1–13.0 37 40 0.52 0.117 0.723
13.1–15.0 81 62 0.43 2.524 0.112
15.1–17.0 114 127 0.53 0.701 0.402
17.1–19.0 114 90 0.44 2.824 0.093
19.1–21.0 207 82 0.28 54.066 0.000
21.1–23.0 180 81 0.31 37.552 0.000
23.1–25.0 79 45 0.36 9.323 0.002
25.1–27.0 43 15 0.26 13.517 0.000
Month
Aug–12 102 49 0.32 18.603 0.000
Sep–12 111 39 0.26 34.560 0.000
Oct–12 40 18 0.31 8.345 0.004
Nov–12 66 8 0.11 45.459 0.000
Dec–12 83 40 0.33 15.033 0.000
Jan–13 109 33 0.23 40.676 0.000
Feb–13 83 45 0.35 11.281 0.001
Mar–13 82 47 0.36 9.496 0.002
Apr–13 42 46 0.52 0.182 0.670
May–13 47 85 0.64 10.939 0.001
Jun–13 32 90 0.74 27.574 0.000
Jul–13 80 69 0.46 0.812 0.368
Fig. 2. Relationship of fecundity with body length (a) and total weight (b)
Total 877 569 0.39 65.604 0.000
of N. furcosus.

426
M.M. Rahman and A.F. Samat
Fig. 3. Monthly mean number (%) of various size oocysts observed in the fe­
male fish ovaries during August 2012–July 2013.
3.4. Development of oocytes
Gonads of female N. furcosus followed six maturity stages: immature,
maturing, mature, ripe and running, spent and resting (Fig. 4). Immature
ovary was observed from August to December with the highest per­
centage (67%) in October (Fig. 5). Immature ovary composed mainly of
oogonia cells (unyolked oocyte) and primary oocytes (atretic oocytes)
(Fig. 4a). Most of the oocytes in the immature ovary were from 0.02 to
0.10 mm in dimeter. The ovaries entered the maturing or developing
stage in November and December during which most of the ovaries
(50–65%, Fig. 5) contained primary oocytes and primary vitellogenic
oocytes (PVO) with yolk granule (Fig. 4b). Oocytes were larger
(0.11–0.30 mm in diameter) and more in number compared to oocytes
427
Aquaculture and Fisheries 6 (2021) 424–431
in the immature stage. A few oogonia were also observed in the
maturing stage. The mature stage of ovaries was observed in January
and February when most of the ovaries (65–70%, Fig. 5) contained
secondary vitellogenic oocyte and post vitellogenic oocyte (Fig. 4c).
These are large and mature oocytes (0.51–0.70 mm in diameter).
Oogonia were also observed scarcely in the mature stage. After the
mature stage, ovaries entered the ripe and running stage during March
to July when most of the ovaries (54–61%, Fig. 5) consisted mainly of
post vitellogenic oocytes consisting a nucleoli at the center of the nu­
cleus (Fig. 4d). At this stage, more yolk accumulation was observed
compared to mature stage. At this stage, most of the oocytes were from
0.91 to 1.20 mm in dimeter. The fish spawn a number of times during
this stage. The spent stage of ovaries was observed in August (Fig. 4e). At
this stage, most of the ovaries (44%) had also oogonia, which were
scattered between secondary vitellogenic oocytes. In addition, ovaries
had large empty spaces where post vitellogenic oocytes were located
before. Finally, the ovaries entered the resting stage in September
(Fig. 4f) during which most of the ovaries (55%, Fig. 5) contained
scattered eggs but not as large as very recently spawned ovaries. Large
space, and few primary oocytes (atretic oocytes; 0.11–0.20 mm in
dimeter) and oogonia (unyolked oocytes; 0.02–0.10 mm in dimeter)
were also observed in the resting stage of ovaries.
3.5. Gonadosomatic index
Monthly mean gonadosomatic index of female and male N. furcosus is
presented in Fig. 7. Female fish had the lowest mean gonadosomatic
index in October (Fig. 6a). A sudden increase of female’s gonadosomatic
index started after December, reaching a peak in March. Gonadosomatic
index then slightly decreased in April and May until another peak in
June. Mean gonadosomatic indexes of female fish in September,
October, November and December were statistically same (P > 0.05)
and were statistically lower compared to remaining sampling months (P
< 0.01). Monthly mean GSIs of male were similar to female with a few
exceptions (Fig. 6b). Male fish did not show any peak GSI in June.
3.6. Sexual maturity
The highest gonadosomatic index was 6.64% in females and 0.97%
in males (Fig. 7a). The gonadosomatic index rose sharply at around and
above 15.7 cm total length in females. Female fish with a GSI ≥2.2%
could approximately be considered as sexually matured females.
Therefore, the SFSM was considered to be around 15.7 cm total length in
females. Male’s GSI increased sharply at around 15.8 cm total length
(Fig. 7b). Therefore, 15.8 cm total length was considered as the SFSM for
male N. furcosus. Male fish with a gonadosomatic index ≥0.28% could
approximately be considered as mature males. The SFSM was calculated
as 15.6 cm total length for females and 15.5 cm total length for male
using the empirical equation developed by Binohlan and Froese (2009).
Using minimum GSI value of mature fish, the number of mature male
and female were calculated as 279 (31.8%) and 215 (37.8%),
respectively.
4. Discussion
This study provides an important contribution towards the primary
literature describing sex ratio, fecundity, relationship of fecundity with
body length and body weight, size of oocyte, oocytes development,
spawning season and sexual maturity of N. furcosus. Especially, by
sampling across a full one-year cycle in N. furcosus fishery and collecting
a plethora of biological data, we have delineated their reproductive
biology. This information can be discussed according to possible life-
history requirements for the N. furcosus and used to not only progress
regional but also broader fisheries management. Analysis of sex ratio
indicated that males dominated in large size fishes. This might be due to
the low survival of female fish and higher life span of male fish (Granada
M.M. Rahman and A.F. Samat Aquaculture and Fisheries 6 (2021) 424–431

Fig. 4. Microscopic observation of various stages of N. furcosus overies. O, oogonia; PO, primary oocyte; PVO, primary vitellogenic oocyte; SVO, secondary vitel­
logenic oocyte; PsVO, post vitellogenic oocyte; ES, empty spaces. Stage I (a); stage II (b); c, Stage III (c); stage IV (d); stage V (e); stage VI (f).

et al., 2004; Wu et al., 2008; Rajkumar, Rahman, Reni Prabha, & Phu­
kan, 2013). In addition, sexual dimorphism and differences in growth
and survival between male and female fish might also be contributing
parameters. In the current observation, we did not observe any sexual
dimorphism in N. furcosus. The growth of male might be greater than
female as the growth of female might be slowed at the start of maturity.
Slowed growth and higher mortality in large size females have to date
been generally accepted possibly due to greater costs associated with
spawning (Farley, Williams, Hoyle, Davies, & Nicol, 2013; Rahman,
2017). Granada et al. (2004) found higher percentage of males than
females in large size N. bathybius at the Kagoshima Bay (Japan) due to a
higher growth rate of male compared to female. The sex ratio of large
size fish may also be related to fecundity (Wu et al., 2008). Some studies
found that for those with high fecundity such as the mullet (Mugil
cephalus), male outnumbered female in large size fish (Chiou & Chen,
1993; Su, 1989). Published data comparing the ratio of male and female
Fig. 5. Percent composition of various reproduction stages observed in female
N. furcosus from the east coast of Malaysian peninsular or elsewhere is
N. furcosus during August 2012–July 2013.
lacking, although a number of studies were reported on other species of
Nemipterus genus from the other geographical regions. For example,

428
M.M. Rahman and A.F. Samat
Fig. 6. Monthly variation of mean (±95% CI) gonadosomatic index (%) in
female (a) and male (b) N. furcosus population during August 2012–July 2013.
Mean with no common letter indicates statistically different (P < 0.05) based on
Tukey test.
Granada et al. (2004) observed the sex ratio of N. bathybius at the
Kagoshima Bay, Japan with males dominating at large-sized fish.
Nemipterus furcosus also showed a temporal variation in sex ratio
with male predominating in most of the sampling months. Almost
similar temporal variation in sex ratio was observed in N. japonicus by
Raje (2002) in Indian coastal water. However, the overall sex ratio
observed in this study is comparable with Persada at al. (2016), who
observed male N. furcosus outnumbered than female N. furcosus at the
Jakarta Bay, Indonesia.
In the present study, the SFSM was estimated using the GSI-TL (total
length) plot and the empirical equation by Binohlan and Froese (2009).
The size at first sexual maturity of fishes can also be obtained from lo­
gistic equation derived from plots of the percentage occurrence of
mature females against length class (Wu et al., 2008). However, many
studies reported (e.g., Hossain, Ahmed, Islam, Jasmine, & Ohtomi,
2010; Hossain & Ohtomi, 2008; Ohtomi et al., 2003) a low accuracy of
the logistic equation in estimating the size of sexual maturity of fishes.
Nowadays, using the GSI-TL plot to determine the size at first sexual
maturity is a common and well-established method. However, in the
present study, both GSI-TL plot and the empirical equation gave almost
an identical result showing the first maturation of male and female
N. furcosus at almost same size (female: 15.7 cm (GSI-TL plot), 15.6 cm
(empirical equation) and male: 15.8 cm (GSI-TL plot), 15.5 cm (empir­
ical equation)). The observed SFSM of male and female N. furcosus
cannot be compared to any other studies as no such study has been found
in the literature. However, Krishnamurthi (1971) determined 16.5 cm
total length as the SFSM of N. japonicus in the Andhra-Orissa coast, India.
Vivekanandan and James (1986) reported lower length (14.5 cm) at first
maturity of N. japonicus in the Madras coastal water, India. Maturity of
fsh can be differed based on the environmental conditions (Jennings,
Kaiser, & Reynolds, 2001; Rahman & Balcombe, 2018; Rahman &
Verdegem, 2007). Water temperature, fishing intensity and abundance
of food are major parameters that influence maturity of fish altough the
information about these variables are not presented in this study.
We recorded that the fecundity of N. furcosus varied from 54,970 to
429
Aquaculture and Fisheries 6 (2021) 424–431
Fig. 7. Relationship of gonadosomatic index (GSI) with total length of (a) fe­
male (n = 569) and (b) male (n = 877) N. furcosus.
236,938 with a mean of 102,477 ± 43,580 (standard deviation). In
comparison, N. furcosus from the Jakarta Bay, Indonesia had a range of
fecundity of only 39,728–40,921 (Persada et al., 2016). In coastal water
of India, a range of fecundity of 10,260–184,946 for N. japonicas was
recorded by Raje (2002), while an even higher range of fecundity of 13,
758–398,859 for N. peronii had been recorded from the waters of
southwestern Taiwan by Wu et al. (2008). Thus, the fecundity of fish is
highly variable depending on many factors including species, size and
maturity stage of fish. However, the documented fecundity of fish may
not be comparable among studies since size and maturity stage of fish
were different. We used matured fish for a full 12-month cycle, while
Persada et al. (2016) used 2–3 years old fish without considering
maturity for a period of only three months. Our observed fecundity was
fluctuated with the length and weight of fish. A significant relationship
was observed for total length and total body weight with fecundity in
N. furcosus. This concurs with many other studies on Nemipterus genus
(Manojkumar, 2004; Nettely et al., 2016; Raje, 2002; Wu et al., 2008).
The reproductive period of fish is commonly determined by their
gonadosomatic index although the study of gonad histology is time
intensive and considered a powerful tool for reproductive studies in fish
as it can precisely determine their spawning season (Kopf, Davie,
Bromhead, & Young, 2012; Zeyl, Oliver, Love, & Higgs, 2014). In the
present study, oocytes diameter, histological examination of ovaries and
monthly mean GSI were used to determine the spawning season. All
these methods gave consistent result about the spawning season of
N. furcosus. Oocytes diameter measurements showed that the largest
sized oocytes 0.91–1.20 mm in diameter were appeared from March to
July. Histological examination of ovaries indicated that the ripe and
running stage of oocytes was appeared during the same period
M.M. Rahman and A.F. Samat
(March–July). Female fish showed comparatively higher GSI during
March to July with two peaks in March and June. Although fish had an
increasing trend of GSI during the December–February period, this
period should not be included in the main spawning season as gonads
were mostly in the maturing stage. Ripe and running stage of gonad was
hardly observed at this period. Fish do not spawn when their gonads are
in the maturing stage because their ovaries contain mostly primary
oocytes (Kokokiris, Stamoulis, Monokrousos, & Doulgeraki, 2014;
Nettely et al., 2016). Based on the above observations, the spawning
season of N. furcosus on the east coast of Malaysian peninsular appears to
be from March to July with peaks in March and June. Similar extended
spawning season is generally observed in other Nemipterus species. For
example, Wu et al. (2008) reported the reproductive period of N. peronii
at southwestern Taiwanese coast from February to July with peaks in
April and May. In another study, Granada et al. (2004) determined the
reproductive period of N. bathybius in Kagoshima Bay, southern Japan
from June to October with a peak-spawning period in July to September.
In conclusion. the result regarding sexual maturity would be helpful
to determine the lowest acceptable harvest length of this species under
the management aim to enter female fish for reproduction minimum one
time in their life. It would also be useful to guide activities that routinely
estimates variations in SFSM associated to various factors including
overfishing (Rahman, 2017). It is necessary to continue the observation
for understanding the yearly maturation variability that associated with
the fish size and the time of the year. Based on this result, N. furcosus
spawners can be protected during March to July to allow all spawners to
spawn minimum one time in their life. The data collected here have
implications for optimal fine− scale harvesting of N. furcosus for their
proper management and also for their artificial breeding. There is
further research potential to examine the reproductive period and SFSM
at various spatial scale to understand the impact of fisheries upon wild
fish populations in various aquatic environments.
CRediT authorship contribution statement
Mustafizur M. Rahman: Methodology, Writing - original draft,
Formal analysis, Data curation. Amira F. Samat: Data curation.
Acknowledgements
We would like to thank the International Islamic University Malaysia
and the Ministry of Higher Education, Malaysia for the financial assis­
tance through P-RIGS18-032-0032 and FRGS19-096-0705 projects,
respectively. No author has any conflict of interest with the content of
the manuscript.
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