Growth, mortality, recruitment pattern, and exploitation rate of shared stock flying fish (Exocoetidae) at border area of Indonesia and Timor Leste in Ombai Strait

Abstract

Flying fish (Exocoetidae) is one important small pelagic fisheries commodity in border areas of Indonesia and Timor Leste. However, the data of population parameters in Ombai Strait are not yet known, which is important to inform unit stock management and sustainable fisheries. This study aimed to provide information about growth and mortality parameters, recruitment pattern, and exploitation rate of flying fish. Data collection was carried out during May 2018-June 2019 by measuring and observing the flying fish landed at the fish landing sites in Belu Indonesia and Bobonaro Timor Leste. Data analysis used the FAO-ICLARM Stock Assessment Tools (FiSAT) program. The results showed that the asymptotic length (L∞) was 330.5 mm, growth constant (K) was 0.99 per year, and growth performance indices (Φ) was 0.50. The natural mortality (M) was 0.92 per year, fishing mortality (F) was 0.12 per year, and total mortality (Z) was 1.04 per year. The highest recruitment occurs in May to September with peak season in June (14.82%) and September (15.16%). Exploitation rate (E) flying fish in the border area of Indonesia and Timor Leste in Ombai Strait is relatively low about 0.11 and still at a rational and sustainable utilization range (E < 0.5).

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Growth, mortality, recruitment pattern, and exploitation rate of shared
stock flying fish (Exocoetidae) at border area of Indonesia and Timor
Leste in Ombai Strait
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International Symposium on Aquatic Sciences and Resources Management
IOP Conf. Series: Earth and Environmental Science 744 (2021) 012062
IOP Publishing
doi:10.1088/1755-1315/744/1/012062
1
Growth, mortality, recruitment pattern, and exploitation rate
of shared stock flying fish (Exocoetidae) at border area of
Indonesia and Timor Leste in Ombai Strait
B M Rehatta1, 3,*, M M Kamal2, M Boer2, A Fahrudin2,5, Zairion2 and J S R
Ninef4
1 Department of Coastal and Marine Resource Management, Graduate School, IPB
University, Bogor Indonesia
2 Department of Aquatic Resources Management, Faculty of Fisheries and Marine
Science, IPB University (Bogor Agricultural University), Bogor Indonesia
3 Faculty of Fisheries and Marine Science, Artha Wacana Christian University,
Kupang, East Nusa Tenggara Indonesia
4 Faculty of Marine and Fisheries, University of Nusa Cendana, Kupang, East Nusa
Tenggara Indonesia
5 Center for Coastal and Marine Resources Studies (CCMRS), IPB University (Bogor
Agricultural University), Kampus IPB Baranangsiang, Jl. Raya Pajajaran No. 1, Bogor
16127, West Java, Indonesia
*Corresponding author: beatrix@ukaw.ac.id
Abstract. Flying fish (Exocoetidae) is one important small pelagic fisheries commodity in
border areas of Indonesia and Timor Leste. However, the data of population parameters in Ombai
Strait are not yet known, which is important to inform unit stock management and sustainable
fisheries. This study aimed to provide information about growth and mortality parameters,
recruitment pattern, and exploitation rate of flying fish. Data collection was carried out during
May 2018-June 2019 by measuring and observing the flying fish landed at the fish landing sites
in Belu Indonesia and Bobonaro Timor Leste. Data analysis used the FAO-ICLARM Stock
Assessment Tools (FiSAT) program. The results showed that the asymptotic length (L∞) was
330.5 mm, growth constant (K) was 0.99 per year, and growth performance indices (Ф) was
0.50. The natural mortality (M) was 0.92 per year, fishing mortality (F) was 0.12 per year, and
total mortality (Z) was 1.04 per year. The highest recruitment occurs in May to September with
peak season in June (14.82%) and September (15.16%). Exploitation rate (E) flying fish in the
border area of Indonesia and Timor Leste in Ombai Strait is relatively low about 0.11 and still at
a rational and sustainable utilization range (E < 0.5).
Keywords: flying fish; Ombai Strait; population dynamic; shared stock
1. Introduction
Flying fish is a small pelagic fish group that has a high economic value because its meat and eggs can
be used [1]. This encourages flying fish to become a target to catch and source of livelihood for
fishermen as well as a source of animal protein for coastal communities. Rehatta et al. [2] stated that
flying fish an important fishery commodity in the border area of Indonesia and Timor Leste.

International Symposium on Aquatic Sciences and Resources Management
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The stock of flying fish in the Ombai Strait is a shared stock used by fishermen in Indonesia and
Timor Leste. Fishermen in Belu (Indonesia) and Bobonaro (Timor Leste) carry out fishing activities in
Ombai strait using drift gillnet. Gillnets dominate the structure of fishing gear in Belu around 49.0% [3]
and Bobonaro around 80% [4]. The production of flying fish in Belu in 2018 was 120.5 tons and
contributed around 22% to the production of small pelagic fisheries [3]. Flying fish production in
Bobonaro District in 2018 was 45.0 tons and contributed around 17% of small pelagic fisheries [4].
Biological aspects of flying fish that have been studied in Indonesia include aspects of reproductive
biology in Tual, Southeast Maluku [5], growth parameters in Binuangeun, Banten [6], species diversity
and size distribution in Eastern Indonesian waters [7], reproductive strategies in Indonesian waters [8],
biology and fisheries of flying fish in the South Makassar Strait [9], maturation and spawning in the
Makassar Strait [10], morphometry, growth factors and conditions in the East Seram Sea [11], species
determination [12]. However, none of these studies have focused on the biology and parameters of the
flying fish population in Ombai Strait.
Given the importance of flying fish resources in Ombai Strait and the lack of data and information
availability, this research is important to provide data and information needed for flying fish sustainable
resources management. One important aspect to support the management of flying fish resources is basic
knowledge about population dynamics, especially parameters of growth, mortality, recruitment, and
exploitation rates aspect.
2. Methods
2.1. Study area
The research was conducted from May 2018 to June 2019. The locations of data collection of fish caught
by fishermen in Belu Regency, Indonesia include Dualaus, Jenilu, Kenebibi, and Silawan villages.
Meanwhile in Bobonaro District of Timor Leste, locations of data collection includes Batugade, Sanirin.
and Aidabaleten villages at Enelaran and Beacou (figure 1).
Figure 1. Research locations in Belu Regency, Indonesia, and Bobonaro District,Timor Leste.

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2.2. Sampling procedure
The collecting of flying fish samples totals 2405 fish from any proportion catch of drift gillnet fisheries
in each month. Sampling was taken continuously every month from May 2018 to June 2019 with
measuring the total length of samples.
2.3. Data analysis
Length frequency analysis was carried out using the FiSAT II (FAO-ICLRAM Stock Assessment) -
ELEFAN (Electronic Length Frequency Analysis) program, which is a program package for analyzing
length-frequency data with the help of a computer device [13]. Estimation of the length at the first mature
(Lm) using the formula, as equation (1) [14]:
 = 0.8979  ∞ − 0.078 (1)
Estimation of the average size first captured (LC) is done by calculating the L50 value. A lethal value
of 50% is considered as the optimum point of fishery resource utilization. The process of estimating the
value of LC is carried out with the FISAT II-ELEFAN software program [15].
Growth parameters were estimated using the von Bertalanffy growth model with mathematical
equation (2) [16]:
  (2)
where Lt is the length of the fish at age t (time unit), L∞ is the theoretical maximum length (asymptotic
length), K is the growth coefficient, t0 is the theoretical age. Estimation of the value of the K and L∞
growth coefficients were carried out using the Ford Walford Plot method which is derived from the von
Bertalanffy model for t equals t + Δt. Estimates for the value t0 (theoretical age of fish when the length
is equal to zero) are obtained through the equation (3) [17]:
       (3)
Estimated maximum fish age (tmax) = 3/K + t0 [17].
The total mortality rate (Z) was estimated using a linear catch curve based on length composition
data [13], which was calculated using the FISAT II program package. The natural mortality rate (M)
was estimated using the empirical formula of Pauly (1980) in [16] as equation (4):
               (4)
M is natural mortality, L∞ is the asymptotic length in the von Bertalanffy growth equation (mm), K is
the growth coefficient in the Von Bertalanffy growth equation, and T is the mean surface temperature
of water (0C). The catch mortality rate (F) is determined by equation (5):
     (5)
The exploitation rate (E) is determined by comparing the fishing mortality rate (F) with the total
mortality rate (Z) [17], as equation (6):
  
   
 (6)
E is the rate of exploitation, M is the natural mortality rate, F is the capture mortality rate and Z is the
total mortality rate.
Recruitment patterns were analyzed with the FISAT II program in the recruitment pattern subprogram
to determine the construction of a time series of recruits from length frequencies in determining the
relative peak recruitment per year.
3. Results
Five species of flying fish were identified (table 1) and length measurements indicated that total length
ranged from 140-314 mm with a mean length of 197 mm (table 2).

International Symposium on Aquatic Sciences and Resources Management
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Table 1. Flying fish species collected from fishermen's catches in Belu and Bobonaro.
No.
Species
Genus
Common name
1
Cheilopogon abei Parin, 1996
Cheilopogon
Abe’s flying fish
2
Cheilopogon intermedius Parin, 1961
Cheilopogon
Intermediate flying fish
3
Cheilopogon spilopterus (Valenciennes, 1847)
Cheilopogon
Many spotted flying fish
4
Cypselurus poecilopterus (Valenciennes, 1847)
Cypselurus
Yellowing flying fish
5
Hirundichthtys oxycephalus (Bleeker, 1852)
Hirundichthtys
Bony flying fish
Table 2. Minimum length (Lmin), maximum length (Lmax), mean length (Lmean), length at first captured
(Lc), and length at first maturity of gonad (Lm) of flying fish in Ombai Strait.
Location
Lmin (mm)
Lmax (mm)
Lmean (mm)
Lc (mm)
Lm (mm)
Belu
140.00
269.00
197.20
-
-
Bobonaro
146.00
314.00
196.70
-
-
Ombai Strait
140.00
314.00
197.03
153.86
152.86
Table 2 shows that the total length of flying fish landed in Belu ranges from 140-269 mm with a
mean length of 197.2 mm, while in Bobonaro ranges in 146-314 mm with a mean length of 196.7 mm.
The average length at first captured (Lc) was 153.86 mm and the average length at first maturity of
gonads (Lm) was 152.86 mm (table 2).
The frequency distribution of the total length of flying fish in Ombai Strait during the study period
was dominated by the size of class 200-214 mm (figure 2). The distribution of the total length-frequency
according to the time of observation is presented in figure 3.
Figure 2. Length frequency distribution of flying fish landed in Belu, Indonesia
and Bobonaro, Timor Leste.
Figure 3 shows that in May-July there are two age groups of fish and another month there is one age
group of fish. In May–July, August–October, and November–January there was no shift mode of length
and the size of the fish structures did not differ significantly. In February, May, and August there is a
shift in the mode of length to the left, and in April and November, there is a shift in mode length to the

International Symposium on Aquatic Sciences and Resources Management
IOP Conf. Series: Earth and Environmental Science 744 (2021) 012062
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doi:10.1088/1755-1315/744/1/012062
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right. November–April the frequency of fish length was dominated by large fish (>197 mm) and in
August, it was dominated by small fish (<197 mm).
Figure 3. Length frequency distribution of flying fish caught in Ombai Strait.

International Symposium on Aquatic Sciences and Resources Management
IOP Conf. Series: Earth and Environmental Science 744 (2021) 012062
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doi:10.1088/1755-1315/744/1/012062
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The analysis of the mode shift in the total length of flying fish based on the time of observation shows
that there are two age groups (cohorts) of flying fish in Ombai Strait (figure 4).
Figure 4. Growth graph of flying fish in Ombai Strait.
The results of the analysis of the growth parameters of flying fish in Ombai Strait show that the
asymptotic length (L∞) is 330.53 mm TL, the growth rate (K) is 0.99 per year, and the age at zero fish
length (t0) is – 0.0832 years. The growth curve of flying fish in Ombai Strait follows the equation Lt =
330.53 (1 - exp [-0.990 (t + 0.0832)]). The maximum age of fish (tmax) is 3.03 years. Growth performance
indices (Ф) was 0.50. The growth curve of flying fish based on the Von Bertalanffy equation in the
waters of the Ombai Strait is presented in figure 5.
Figure 5. Von Bertalanffy growth graph of flying fish in Ombai Strait.
A linear analysis of fishing yield curves to estimate the total mortality rate (Z) of flying fish in Ombai
Strait gives a yield of 1.039 per year (figure 6). The natural mortality rate (M) at 290C is 0.922 per year
and the fishing mortality rate (F) is 0.117 per year. The estimated exploitation rate (E) of flying fish in
Ombai Strait is 0.11.

International Symposium on Aquatic Sciences and Resources Management
IOP Conf. Series: Earth and Environmental Science 744 (2021) 012062
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doi:10.1088/1755-1315/744/1/012062
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Figure 6. Length-converted catch curve of flying fish in Ombai Strait.
Pattern recruitment of flying fish in Ombai Strait shows that recruitment occurs throughout the year.
The highest recruitment occurred in the period May-September with peak recruitment occurring in
September and June with relative recruitment values of 15.16% and 14.82% (figure 7).
Figure 7. Recruitment pattern graph of flying fish in Ombai strait.

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4. Discussion
The length range of flying fish in the border areas of Indonesia and Timor Leste in Ombai Strait is 140
- 314 mm TL with a mean length of 197.0 mm TL. The length range is wider and has a maximum length
(Lmax) that is greater than the size of the flying fish as a result of previous studies. Harahap and Djamali
[6] stated that the length of flying fish (Hirundichthys oxycephalus) in Binuangeun Banten waters ranges
from 214.5-278.5 mm TL. Furthermore, Tuapetel et al. [11] reported that the length range of flying fish
in the East Seram Sea was 208.7-274.4 mm FL. The difference in the length range is thought to be due
to differences in aquatic environmental conditions [18] and differences in the fishing gear used [19, 11].
The lengths of flying fish caught in Ombai strait and landed in Belu (Indonesia) and Bobonaro (Timor
Leste) indicate differences in the total length range. The flying fish landed at Belu had a smaller total
length range (140-269 mm) than the flying fish landed at Bobonaro (146-314 mm). Differences in
geographic location [20] and the conditions of the aquatic environment [18] may have influenced the
differences in length. Ali [1] also stated that the overfishing factor can lead to population diversity
between flying fish groups. Furthermore, Harahap and Djamali [6] stated that the level of exploitation
can affect the total length range of flying fish.
The estimate of the length at first captured (Lc) of flying fish in Ombai strait of 153.86 mm TL is
relatively the same as the length at first captured of flying fish Cypselurus oxycephalus. C. oligolepis,
and Exocoetus volitans in Makassar Strait, ranging from 150-160 mm [9]. Dalzell [21] reported Lc of
flying fish Cheilopogon nigrican was 149 mm, Cypselurus opisthopus was 140 mm, and
Oxyporhamphus convexus was 152 mm in Camotes Sea Philippines. The difference in Lc size between
fish species and populations is influenced by factors of differences in fishing gear and methods and fish
size distribution [22].
The length at first maturity of gonads (Lm) of flying fish in the border area of Indonesia and Timor
Leste in Ombai Strait is 152.86 mm TL. These results indicate a slight difference with Lm Hirundichthys
oxycephalus in the Makassar Strait of 151.1 mm FL [23], male Cypselurus oxycephalus 132.6 mm and
female 117.7 mm, male Cypselurus oligolepis 145.7 mm and female 120.8 mm, male Exsocoetus
volitants 114.2 mm and 138.2 mm [9], Cypselurus oligolepis 229.0 mm and C. spilopterus 284.8 mm in
Tual [5]. Oliveira et al. [24] stated that the size at the first maturity of gonads is not the same and even
varies between individuals of the same species. Furthermore, Ndiaye et al. [25] stated that the length at
the first maturity of gonads varies according to sex, site, and season. This difference is influenced by
the availability of energy for gonad development [26], differences in fishing grounds, fishing pressure,
sampling time and season [27], and length composition [28].
Estimation of the length at first captured can be used as a reference in determining fisheries resource
management efforts [29]. Further explanation that if the fish has a size of Lc> Lm, then this situation
does not endanger the fish population because the fish are allowed to spawn. The length at the first
maturity of gonads is an important parameter in determining the size of the mesh and the minimum size
of fish that can be caught [28, 24]. The results of this study indicate that the length at first captured is
greater than the length at first maturity of the gonad (Lc>Lm). Thus, the use of gillnets with a mesh size
as currently used by fishermen in Belu and Bobonaro can still be maintained at a mesh size of 1.25
inches, and does not endanger the flying fish population in Ombai Strait because the fish caught are
dominated by adult fish.
The estimated asymptotic length (L∞) of flying fish in Ombai Strait of 330.5 mm TL tends to be
larger than that which has been reported. Dalzell [21] reported that L∞ Cheilopogon nigricans and
Cypselurus opisthopus in the Comotes Sea Philippines were 210 mm and 280 mm, respectively. Further
reported by [30] that the asymptotic length of Hirundichthys affinis in the Barbados Sea was 245 mm,
while Hirundichthys oxycephalus was 254 mm in the East Taiwan Sea [31]. In Indonesia, several
researchers have reported the asymptotic length of the flying fish Hirundichthys oxycephalus was 182
mm in the Flores Sea and Makassar Strait [1], 321.1 mm in Binuangeun Banten [6], 245 mm in East
Seram Sea [11]. The growth coefficient (K) of flying fish in Ombai strait was 0.99 per year, which is
greater than the growth coefficient of Hirundichthys oxycephalus in Binuangeun Banten [6] and in the
East Taiwan Sea [30], but smaller than the growth coefficient of Cheilopogon nigricans and Cypselurus

International Symposium on Aquatic Sciences and Resources Management
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doi:10.1088/1755-1315/744/1/012062
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opisthopus in the Comotes Sea, Philippines [21], Hirundichthys affinis in the Barbados Sea [31] and
Hirundichthys oxycephalus in the Flores Sea and Makassar Strait [1], and the East Seram Sea [11] (table
4). This result indicates that the flying fish in Ombai Strait are estimated to be able to grow to a maximum
length of 330.5 mm TL with a growth rate of 0.99 per year and a maximum age (tmax) of 3.03 years. [32]
stated that the difference in the value of the growth coefficient (K) and the asymptotic length (L∞) is
probably caused by the structure of the data collected and data analysis performed. Furthermore,
Motlagh et al. [18] stated that this difference is thought to be due to variations in fishing intensity and
environmental conditions. The maximum length (Lmax) of flying fish caught from the Ombai Strait is
314 mm. This shows that the fishing intensity is still low and the waters are relatively fertile so that the
flying fish population can live lengthier and the size is close to the asymptotic length.
Table 3. Growth parameter of flying fish at various locations in Indonesia and surrounding waters.
Site
Species
Asymptotic
length (L∞)
(mm)
Growth
coefficient (K)
(per year)
Reference
Camotes Sea, Philippines
Cheilopogon nigricans
210.0
1.75
[21]
Cypselurus opisthopus
280.0
2.05
Barbados Sea
Hirundichthys affinis
245.0
2.90
[31]
Flores Sea, Makassar Strait,
Indonesia
Hirundichthys oxycephalus
182.0
1.30
[1]
Binuangeun, Banten Indonesia
Hirundichthys oxycephalus
321.1
0.15
[6]
Taiwan East Sea
Hirundichthys oxycephalus
254.0
2.60
[30]
East Seram Sea, Indonesia
Hirundichthys oxycephalus
245.0
1.80
[11]
Ombai Strait, Indonesia
Cheilopogon abei
C. spilopterus
C. intermedius
Cypselurus poecilopterus
Hirundichthys oxycephalus
330.5
0.99
This study
The total mortality rate (Z) of flying fish in Ombai Strait is 1.039 per year and is lower than the total
mortality of Cheilopogon nigricans and Cypselurus opisthopus in the Camotes Sea Philippines of 5.44
per year and 10.47 per year as reported by Dalzell [21]. The same thing happened to the natural mortality
rate (M=0.922 per year) and fishing mortality (F=0.117 per year) which were lower than the other
locations. The natural mortality rate (M) is higher than the catch mortality rate (F) of flying fish in
Ombai Strait. This indicates that the flying fish population experiences high levels of predation and
competition. resulting in high natural mortality as well. [16] stated that natural mortality occurs due to
various reasons including predation, disease, spawning stress, hunger, and old age. Furthermore, Morgan
[26] stated that the different mortality rates between waters are caused by fishing gear targets. Migration,
and ontogenic factors. In this study, the value of the Z/K ratio was 1.05 and less than 2, which means
that the growth of flying fish in the Ombai Strait is more dominant than mortality, so even though the
growth coefficient is relatively not high, the maximum length that can be achieved is high.
The pattern of flying fish recruitment in Ombai strait occurs throughout the year with the highest
recruitment occurring in the period from May to September with peak recruitment occurring in
September and June. Factors that influence recruitment include the availability of adult stocks,
reproductive success, pre-recruitment mortality both at the larval and juvenile stages [33]. Furthermore,
Nugroho and Chodrijah [34] stated that the size of the recruits is determined by the number of brood
stock ready to spawn and the mortality in the period between spawning and fish reaching the stock size.
Besides, the success of recruitment is also determined by the environmental conditions during the post-
larval cohort on the nursery ground. The population of flying fish in Ombai Strait is dominated by adult-

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sized fish (>Lm) and the mortality rate is low, resulting in high availability of brood stock fish that are
ready to spawn. For this reason, to keep flying fish recruits in Ombai Strait high. it is necessary to
regulate fishing activities so that more adult fish are available and ready to spawn.
The utilization rate of flying fish (E) in Ombai strait is 0.11 per year, lower than that of the Camotes
Sea Philippines which ranges from 0.45 to 0.75 per year [21]. The E value of 0.11 per year indicates
that the utilization rate of flying fish in Ombai strait is below the optimum conditions. Pauly [35] stated
that the value of rational and sustainable utilization rates in the range of E<0.5 or the highest is E=0.5.
The utilization rate of flying fish in Ombai Strait is low and can still be increased to reach an optimum
utilization rate of 0.5. This can be done by increasing the fishing mortality rate through an increased
fishing effort by fishermen in Belu and Bobonaro.
5. Conclusion
Five species of flying fish (Exocoetidae) were found in Ombai Strait and adult fish groups (>52.8 mm
TL) dominated the flying fish population. Growth is classified as fast with a growth rate of 0.99 per year
and an asymptotic length of 330.5 mm TL which is achieved at the age of >3 years. The natural mortality
rate is higher than the fishing mortality rate of flying fish in Ombai Strait. The pattern recruitment of
flying fish in Ombai Strait occurs throughout the year with peak recruitment occurring in September
and June. The exploitation rate of flying fish in Ombai Strait is still within the limits of rational and
sustainable use. The conclusion of this study provides information related to biological aspects that can
be used as a basis for making flying fish fisheries management policies in the border areas of Indonesia
and Timor Leste in Ombai Strait.
Acknowledgments
The author thanks to the Ministry of Research, Technology, and Higher Education through the
Directorate General of Higher Education for funding research funds through BPPDN. We also thank to
the Diração Nacional de Pesca e Aquicultura, Ministério de Agricultura e Pescas of República
Democrática de Timor Leste, Fisheries Agency of Belu, and all those who helped this research in Belu
and Bobonaro.
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