Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023)

Original article DOI: 10.19027/jai.22.2.179-186

The ontogenic study of early life stages of culture-bred Nomorhamphus sp.
(Zenarchopteridae) from Lindu, Central Sulawesi

Studi ontogeni stadia awal hidup pada budidaya Nomorhamphus sp.

(Zenarchopteridae) asal Lindu, Sulawesi Tengah

Muh. Herjayanto1,2, Odang Carman1, Dinar Tri Soelistyowati1*, Alimuddin1, Aryo

Wenang Wicaksono1, Harton Arfah1
1
Department of Aquaculture, Faculty of Fisheries and Marine Sciences, IPB University, Bogor, West Java 16680,
Indonesia
2
Department of Fisheries Science, Faculty of Agriculture, University of Sultan Ageng Tirtayasa, Serang, Banten,
Indonesia
*Corresponding author: dinar@apps.ipb.ac.id

(Received March 8, 2023; Accepted May 29, 2023)

ABSTRACT

Nomorhamphus sp. is a freshwater fish that has been traded as an ornamental fish. This fish is unique as an
endemic species with a halfbeak-like jaw and orange color on the caudal fin. However, this fish culture information
needs a further information. Based on this condition, it is necessary to conduct a study as a basis for ornamental
fish breeding and growing-out activities through domestication. A crucial problem in this fish is larval rearing,
which can be observed through ontogeny studies. The study was conducted on the newly-born larval behavior,
morphological development, andropodium development, growth, and survival rate at the early stages, namely
larvae to juvenile. The results showed that the newly-born larvae of Nomorhamphus sp. Lindu had a total length
of 1.6-1.8 cm. Larvae could swim four hours 22 minutes after birth and feed artemia nauplii with surface feeding
type. The initial juvenile stage occurred 25 days of post-birth period with a total length of 2.0-2.2 cm. The water
condition of the rearing during the study could support the larval transformation to juvenile. This study is the first
report related to the aquaculture success of the early life stage of Nomorhamphus sp. Lindu at the domestication
stage.

Keywords: andropodium, domestication, endemic halfbeak, larva development, surface feeding

ABSTRAK

Nomorhamphus sp. adalah ikan air tawar yang telah diperdagangkan sebagai ikan hias. Ikan ini memiliki keunikan
pada statusnya sebagai spesies endemik, bentuk mulut menyerupai paruh setengah (halfbeak), dan warna oranye
pada sirip ekor. Namun informasi budidayanya belum diketahui dengan baik. Karena itu perlu dilakukan kajian
sebagai dasar dalam kegiatan pengembangbiakan dan pembesaran sebagai ikan hias melalui domestikasi. Salah
satu kegiatan penting dalam budidaya yaitu pemeliharaan larva yang dapat diamati melalui studi ontogeni. Kajian
pada studi awal ini dilakukan pada stadia awal hidup yaitu larva sampai juvenil. Pengamatan dilakukan pada tingkah
laku larva pascalahir, perkembangan morfologi, perkembangan andropodium, pertumbuhan dan sintasan pada
lingkungan budidaya. Hasil penelitian menunjukkan bahwa larva Nomorhamphus sp. Lindu yang baru dilahirkan
memiliki panjang total 1,6-1,8 cm. Larva telah dapat berenang pada umur empat jam 22 menit pascalahir (pcl) dan
bisa makan naupli artemia dengan tipe surface feeding. Stadia awal juvenil terlihat pada umur 25 hari pcl dengan
ukuran panjang total 2,0-2,2 cm. Kondisi media pemeliharaan selama penelitian dapat mendukung kehidupan
larva sampai juvenil. Penelitian ini merupakan catatan pertama terkait keberhasilan budidaya stadia awal hidup
Nomorhamphus sp. Lindu pada tahap domestikasi.

Kata kunci: andropodium, domestikasi, ikan endemik, tipe makan permukaan, perkembangan larva
180 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023)

INTRODUCTION ontogenic early life period of Nomorhamphus sp.

Lindu in the cultural environment. The ontogenic
Nomorhamphus is a tropical freshwater genus studies are mainly focused on the initial larval
from the Zenarchopteridae family. Seventy development to become juvenile. Observations
percent of Nomorhamphus species are found as were performed following the post-birth larval
endemic species in Sulawesi Island (Huylebrouck behavior (pbl), morphological development,
et al., 2014; Hadiaty, 2018; Kraemer et al., 2019a; andropodium development, growth, and survival
Kobayashi et al., 2020). The habitats of this fish rate in the culture environment.
are freshwater rivers and lakes (Herjayanto et al.,
2019; Kraemer et al., 2019a; Kobayashi et al., MATERIALS AND METHODS
2020), as reported in Lindu Lake, Central Sulawesi
(Wicaksono et al., 2022). Morphologically, this Fish Catching
fish has a distinct superior jaw, as the lower jaw Fish samples were caught in the inlet flow
is longer than the upper jaw, called a halfbeak of Lindu Lake at >1,000 asl (1°22’19.20” S
(Kraemer et al., 2019a; Kobayashi et al., 2020). 120°01’40.80” E). using a net at 5.5-mm mesh
This fish also has a unique reproduction size. The total female fish caught was nine
strategy through internal fertilization (viviparous), individuals with a total length of 6.6-9.0 cm.
thus male fish have an additional sexual organ After being caught, the fish were maintained by
called andropodium. This organ functions as a stocking them in an aerated container. During
spermatozoan distributor to female genital pore maintenance, the fish remained unfed.
during the mating process (Kraemer et al., 2019b). Maintenance was performed for 24 hours. Fish
The uniqueness of endemic species causes were packed and transported in a closed system,
Nomorhamphus sp. is collected by hobbyists or following Hadiroseyani et al. (2016), Herjayanto
foreign ornamental fish collectors. Moreover, the et al. (2018), and Herjayanto et al. (2020), which
color and shape are also unique as part of the were modified at low stocking density. The initial
aquascape biota. However, these fish are supplied transportation was performed from Lindu Lake
from natural catch, but limited culture studies to Palu City through a land route. Then, fish
information. were sent through an air route from Palu City to
Wicaksono et al. (2022) have reported Soekarno-Hatta Airport, followed by a land route
the initial characterization of biometrical and from the airport to Laboratory of Aquaculture,
morphological approaches of Nomorhamphus sp. University of Sultan Ageng Tirtayasa, Banten. In
from Lindu. Yet, information regarding breeding, this location, fish were reared in a controlled tank.
larval rearing, and controlled seed maintenance
remains unknown. In general, studies that have Fish acclimatization
been conducted in Nomorhamphus species in Acclimatization was performed based on
Central Sulawesi are species description (Kraemer the wild fish maintenance method modified by
et al., 2019a) and the latest habitat condition Herjayanto et al. (2018) and Hadiroseyani et al.
(Bandjolu et al., 2021; Kraemer et al., 2019a). (2016) by stocking the fish in a packing plastic in
Therefore, a cultural approach for Nomorhamphus the rearing tank for 30 minutes for temperature
sp. rearing is necessary to support its sustainable adaptation. Then the fish was removed from the
production as an ornamental fish. Early life history plastic and allowed to escape slowly into the
observation is highly important to gain basic data rearing tank for acclimatization usingaquarium of
for larval rearing management (Dos Santos et al., 100×50×40-cm3.
2016).
This observation is related to morphological Handling Pregnant-broodstock
development, namely body shape and behavior The pregnant female broodstock of
during the developmental stage, which can be Nomorhamphus sp. from the wild were separated
called an ontogenic study (Baras et al., 2012; and reared solitary (Figure 1). The rearing tank
Sørensen et al., 2016; Tinguely et al., 2019). The for pregnant female fish used an aerated clear
ontogenic study in fish is necessary to provide plastic container at 35 cm×25 cm×15.5 cm size.
information for further breeding activities in the The total pregnant female fish was five individuals
nursery sector as part of the aquaculture activities with a total length of 7-8 cm. During the rearing,
(Baras et al., 2012). Based on the following the female broodstocks were fed with live feeds,
condition, a study was performed to describe the namely Tubifex sp. and Daphnia sp. three times
 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023) 181

a day with ad libitum method. Broodstocks were with water at 10 cm height. During the rearing,
reared until giving birth to the larvae. water quality parameters, namely temperature,
dissolved oxygen, and pH were measured. Feed
Larval rearing residue and fecal waste were siphonized once in
The 40 individuals of Nomorhamphus sp. three days.
larvae were reared in an aerated clear plastic
container at 35 cm×25 cm×15.5 cm size. The Data analysis
rearing tank was filled with water until 5 L Data from behavior, ontogenic morphology,
volume. Larval rearing used five larvae that growth, i.e., total length, head length, lower jaw
were stocked at 1 larvae/L. Larvae were fed with length, and anal fin length (Figure 2), and survival
Artemia nauplii at one day of post-birth (pbl). rate were analyzed descriptively using Microsoft
Rearing was performed for 30 days after giving Excel 2010. The analysis results are presented in
birth. Behavior, ontogenic morphology, growth figures and tables.
(total length, head length, lower jaw length, and
fin length), and survival rate were observed at 0, RESULTS AND DISCUSSION
15, 25, and 30 days of pbl.
The larval swimming behavior was observed Results
from the 10 post-birth larvae. The ontogenic The newborn larvae of Nomorhamphus sp.
morphology development was documented in a had a total length of 1.6-1.8 cm (1.69 ± 0.08
photo to observe the difference between larval and cm) (n=3). Larvae could open their mouth with
juvenile stadia, as modified by Baras et al. (2012) an orange-colored lower jaw. The pectoral,
and Wicaksono et al. (2022). The photo was taken ventral, anal, dorsal, and caudal fins have become
in a glass container at 24 cm×7 cm×15 cm, filled definitive organs (Figure 3). The newborn larvae

Figure 1. The pregnant female broodstock morphology of Nomorhamphus sp. Lindu.

Figure 2. Larval measurement method in 0 days of post-birth (above) and juvenile at 25 days of post-birth (below)
in Nomorhamphus sp. Lindu. a. Total length; b. Head length; c. Lower jaw length; d. Anal fin length. Line scale:
0.5 cm.
182 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023)

existed on the bottom contained and occasionally the body pigmentation started to spread on the
swam onto the water surface. transparent body surface (Figure 4).
Based on the behavior of 10 larvae, 60% of The digestive tract of Nomorhamphus sp.
the larvae could swim on the water column at 20 Lindu larvae were still visible until 15 days of pbl
minutes of post-birth (pbl), then increased by 90% as the belly was still transparent. After 20 days
at 3 hours 6 minutes of pbl. All larvae could swim of pbl, the fish belly became non-transparent, as
after 4 hours 22 minutes of pbl. This swimming the Artemia consumed by the larvae was invisible
capability has occurred as the swimming bladder (Figure 5). The anal fin as a secondary sexual
has been filled. The swimming position of the character in male fish (andropodium) was yet
larvae at early life stage was only swimming with visible at 15 and 20 days of pbl. The orange color
their head and water surface in perpendicular in the ventral fin and caudal fin edge was visible
position. when the fish reached 25 days of pbl.
As the larvae grew older, they could swim onto The anal fin of Nomorhamphus sp. Lindu,
the water column horizontally. After horizontal which was modified as andropodium began to
swimming, larvae could chase the Artemia emerge at 25 days of pbl (Figure 6). Therefore, the
nauplii. During the developmental stage, the early stage of juvenile stadia in Nomorhamphus
Nomorhamphus sp. larvae had a morphological sp. Lindu emerges at 25 days of pbl. The total
change in their belly shape. Based on the length of Nomorhamphus sp. juvenile was 2.11
observational results in five days of pbl, larvae cm with a head length of 0.47 cm, lower jaw
had a digestive tract full of Artemia nauplii. Also, length of 0.11 cm, and anal fin length of 0.30 cm
(Table 1).

Figure 3. Ontogenic morphology of the newly-born Nomorhamphus sp. Lindu larvae. Fish size is presented in
Table 1.

Figure 4. Ontogenic morphology of Nomorhamphus sp. Lindu larvae. The newborn larvae (above) and five days of
pbl larvae (center and below). Fish size is presented in Table 1.
 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023) 183

The survival rate of Nomorhamphus sp. at the the larval and juvenile life of Nomorhamphus
final rearing period (30 days of pbl) was 75-100% sp. Based on the water quality parameters
(90.60 ± 11%). Fish death has only occurred at measurement, the temperature was maintained
three days of pbl and 11 days of pbl. The water at 22.30-25.50°C (24.00 ± 0.62°C), dissolved
quality during the rearing period could support oxygen at 4.80-8.90 mg/L (6.85 ± 1.19 mg/L),
and pH at 7.19-9.41 (8.08 ± 0.58).

Figure 5. Ontogenic morphology of Nomorhamphus sp. Lindu larvae at 15 days of pbl (above) and 20 days of pbl
(below). Fish size is presented in Table 1.

Figure 6. Ontogenic morphology of Nomorhamphus sp. Lindu juvenile at 25 days of pbl. Male (above) and female
(below). Fish size is presented in Table 1.

Table 1. Survival rate (n=40) and growth (n=12) of Nomorhamphus sp. Lindu. early life stages.
Age (days of post-birth)
Parameter
0 15 25 30
Developmental stage Larvae Larvae Juvenile Juvenile
Survival rate (%) 100.00 ± 00 90.60 ± 11 90.60 ± 11 90.60 ± 11
Total length (cm) 1.69 ± 0.08 2.10 ± 0.05 2.11 ± 0.08 2.69 ± 0.19
Head length (cm) 0.32 ± 0.04 0.45 ± 0.03 0.47 ± 0.03 0.53 ± 0.09
Lower jaw length (cm) 0.08 ± 0.02 0.10 ± 0.00 0.11 ± 0.01 0.14 ± 0.02
Anal fin length (cm) 0.24 ± 0.03 0.29 ± 0.01 0.30 ± 0.00 0.39 ± 0.05
184 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023)

Discussion (initial juvenile stage), the orange color in the

The early life stage of fish requires more ventral fin and caudal fin edge was visible.
thorough management as larvae are susceptible Fish sexual dichromatics and dimorphism can
to environmental changes and feed availability. be used as secondary sexual characteristics in
Therefore, an ontogenic study of early life stages fish (Wicaksono et al., 2022). In viviparous fish,
is important to provide basic data for larval rearing Xenotoca eiseni, orange color in male fish caudal
management (Baras et al., 2012; dos Santos et al., fin initially emerge at four weeks of pbl or 28
2016). It has been confirmed that Nomorhamphus days of pbl (Tinguely et al., 2019). The secondary
sp. is viviparous fish, as embryonic development sexual character observed in Nomorhamphus
occurs in the female gonad and birth larvae sp. is andropodium in male fish (Kraemer et al.,
(Kraemer et al., 2019a; Kraemer et al., 2019b; 2019b). In this study, andropodium was confirmed
Kobayashi et al., 2020). Based on the newborn to emerge at 25 days of pbl. Thus, 25 days of pbl
Nomorhamphus sp. Lindu larvae observation, is considered as the initial stage of juvenile in
larvae were unable to swim and remained in the Nomorhamphus sp. Lindu.
bottom container. The juvenile fish can grow as a young fish then
This condition has occurred as Nomorhamphus become a mature broodstock. It has been reported
sp. Lindu fish are lecithotrophic viviparous fish, that sexual maturity in another viviparous fish,
as their larvae have yolk as a nutrient source namely P. reticulata, is occurred at 8-10 weeks
during the developmental stage in the female of pbl or 56-70 days of pbl, then becomes a
gonad. This reproductive strategy causes the broodstock at six months of pbl or around 180
newborn Nomorhamphus sp. larvae to require days of pbl (Shahjahan et al., 2013). The sexual
more time to develop and swim freely by counting dimorphism in the anal fin, namely gonopodium,
on the yolk energy remaining after birth. Another is visible at 28 days of pbl (Ahasan et al., 2014).
lecithotrophic viviparous fish is Poecilia vivipara The modified anal fin in Xenotoca eiseni was
(Poecillidae), which has yolk as a nutrient source initially emerged at four weeks of pbl or 28 days
until the final stage of embryonic development of pbl (Tinguely et al., 2019). The secondary
(Arcanjo et al., 2014). During larval development, sexual character formation in modified anal fin
transformation emerges in the organ system and was affected by the androgen hormone, based
body shape (Shahjahan et al., 2013; Herjayanto on the expression of sonic hedgehog (Shh) gene
et al., 2017). The Nomorhamphus sp. larvae (Ogino et al., 2004).
gradually performed a jerky motion. The live feed, Artemia nauplii, can be used
This behavior is an effort for larvae to fill as a feed for Nomorhamphus sp. Lindu larvae, as
the swimming bladder (Herjayanto et al., 2017). this feed is suitable with larval mouth opening.
Another viviparous species, P. reticulata, has It has been reported that Artemia nauplii had a
newborn larvae that can swim, feed, and move size of 379-480 μm, commonly used as a live
away from danger (Shahjahan et al., 2013). In feed for fish culture (Piotrowska et al., 2021).
rainbow fish larvae, Iriatherina werneri, the The newborn Nomorhamphus sp. Lindu larvae
newborn larvae spend most of their time resting were incapable of feeding Daphnia due to its
in the bottom rearing tank, while the fish in the size of 0.48-0.81 mm (Ebert, 2022). However,
juvenile stage can swim in the water column. this study did not measure the size of the larval
Swimming activity elevates as age increases mouth opening, so it is necessary to carry out a
(Herjayanto et al., 2017). After the swimming more detailed characterization of the ontogeny of
bladder has been filled, the Nomorhamphus sp. the larval mouth opening in future studies. The
larvae could swim well to chase Artemia nauplii preliminary observations indicate that Daphnia
as their feed. can be applied at 30 days of pcl.
This external feeding becomes an energy Another halfbeak species, Zenarchopterus
source for larvae to conduct metabolism, grow, dunckeri, has been reported to change its diet as it
and develop. The larval body shape causes grows in nature. In general, juvenile fish feed more
changes and development to reach a definitive on zooplankton and small gastropods, while larger
shape along with increasing age. As they were juveniles feed land insects, such as ants and flies
newly born, the Nomorhamphus sp. larvae had (Kanai et al., 2017). In another halfbeak species,
a transparent body. The morphological change Hemiramphus far, is known as an omnivorous
started to emerge at 20 days of pbl, as the belly fish, as it feeds on fish, isopods, ostracods,
started to become transparent. At 25 days of pbl Daphnia, shellfish, algae, and unidentified parts
 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023) 185

of plants and animals (Tabassum et al., 2017). vivipara (Cyprinodontiformes: Poeciliidae).

The superior jaw shape of the Nomorhamphus, Acta Zoologica 95: 493–500.
where the lower jaw is longer than the upper Bandjolu KP, Madiyono E, Herjayanto M, Gani A,
jaw, indicates that this fish is a type of surface Nur M, Laheng S, Gundo MT. 2021. Checklist
feeding, as seen in Z. dunckeri (Hirota et al., of endemic (Adrianichthyidae, Gobiidae,
2015), H. far (Tabassum et al., 2017), and other Zenarchopteridae) and introduced fish in Lake
Nomorhamphus as endemic species in Sulawesi Poso, Sulawesi, Indonesia. IOP Conference
(Kraemer et al., 2019a; Kobayashi et al., 2020). Series: Earth Environmental Science 869:
The success of maintaining the early life stages in 012060.
this study provides important information for the Baras E, Slembrouck J, Priyadi A, Satyani D,
culture management of Nomorhamphus sp. Lindu Pouyaud L, Legendre M. 2012. Biology and
is an ornamental fish. culture of the clown loach Chromobotia
macracanthus (Cypriniformes, Cobitidae):
CONCLUSION 3- Ontogeny, ecological and aquacultural
implications. Aquatic Living Resources 25:
The Nomorhamphus sp. Lindu larvae have 119–130.
a total length of 1.6-1.8 cm and can swim after Dos Santos JE, Sales NG, dos Santos ML, Arantes
4 hours and 22 minutes post-birth (pbl). These FP, Godinho HP. 2016. Early larvae ontogeny
larvae feed on Artemia nauplii through the surface of the neotropical fishes: Prochilodus
feeding method. The juvenile stage has occurred costatus and P. Argenteus (Characiformes:
at 25 days of pbl with a total length of 2.0-2.2 cm. Prochilodontidae). Revista de Biología
The rearing media condition can support the early Tropical 64: 537–546.
life stages of Nomorhamphus sp. Lindu. Ebert D. 2022. Daphnia as a versatile model
system in ecology and evolution. EvoDevo 13:
ACKNOWLEDGMENTS 1–13.
Hadiaty RK. 2018. Taxonomical status of
We thank the Directorate of Higher Educations endemic freshwater ichthyofauna of Sulawesi.
for funding this research through the PDUPT Indonesian Journal of Ichthyology 18: 175–
2020-2022 scheme with contract number: 1/ 190.
E1/KP.PTNBH/2021 LPPM IPB University. Hadiroseyani Y, Sukenda S, Surawidjaja EH,
The authors would also like to thank Edo and Utomo NBP, Affandi R. 2016. Survival rate of
assistants in the Laboratory of Fisheries Science transported ricefield eels, Monopterus albus
Study Program, Faculty of Agriculture, University (Synbranchidae), in open and closed system
of Sultan Ageng Tirtayasa, for their assistance at water salinity level of 0 and 9 g L-1. AACL
and for providing the research facilities. Also, Bioflux 9: 759–767.
we would like to thank the Sulawesi halfbeak Herjayanto M, Carman O, Soelistyowati DT.
research team, Kamal, Arya, Riva, Dinda, the 2017. Embryogenesis, larval development
ERA team (Agustino, Ando, Fadli, Waris, Adan, and reproduction viability of Threadfin
and Adi), and all field and laboratory teams who Rainbowfish Iritherina werneri Meinken,
contributed to the research. This research is also 1974 Iriatherina werneri Meinken, 1974
presented to Professor Muhammad Zairin Junior under laboratory conditions. Jurnal Akuatika
for his direction and guidance in this research. Indonesia 2: 1–10.
Herjayanto M, Gani A, Adel YS, Suhendra N.
REFERENCES 2019. Freshwater fish of lakes and it’s inlet
rivers in Sulawesi Tengah, Indonesia. Journal
Ahasan MM, Rahman MA, Shahjahan RM, of Aquatropica Asia 4: 1–9.
Begum RA, Alam MM, Habiba SU. 2014. Herjayanto M, Syamsunarno MB, Prasetyo NA,
Studies on larval development of mosquitofish, Mauliddina AN, Agung LA, Widiyawan ER,
Gambusia affnis (Baird and Girard, 1853) in Rahmawati N, Irianingrum N, Nurkhotimah
Bangladesh. Academic Journal of Entomology E, Gani A, Salsabila VN. 2020. Preliminary
7: 109–114. study of closed system transportation, rearing
Arcanjo RB, de Souza LP, Rezende CF, Silva and observation of the eggs Oryzias javanicus
JRF. 2014. Embryonic development and (Bleeker 1854) from Tunda Island. Jurnal
nourishment in the viviparous fish Poecilia Iktiologi Indonesia 20: 93–103.
186 Muh. Herjayanto et al. / Jurnal Akuakultur Indonesia 22 (2), 179–186 (2023)

Herjayanto M, Waris A, Suwarni Y, Halia M, dependent development of a modified anal

Gani A, Findayani N, Cahyani R. 2018. Study fin, gonopodium, as a model to understand
habitat and closed system transportation the mechanism of secondary sexual character
of Sarasin’s Buntingi Oryzias sarasinorum expression in vertebrates. FEBS Letters 575:
Popta, 1905 endemic Lake Lindu as the basis 119–126.
for domestication. Jurnal Akuatika Indonesia Piotrowska I, Szczepkowska B, Kozɫowski
3: 103–109. M. 2021. Influence of the size and form of
Hirota K, Asaoka R, Nakae M, Sasaki K. 2015. Artemia sp. nauplii on the growth and survival
Zenarchopterus dunckeri (Beloniformes: of Atlantic sturgeon (Acipenser oxyrinchus
Exocoetoidei: Zenarchopteridae): an example Mitchill) larvae. Fisheries & Aquatic Life 29:
of adaptation to surface feeding in fishes. 69–79.
Ichthyological Research 62: 286–292. Shahjahan RM, Ahmed MJ, Begum RA, Rashid
Huylebrouck J, Hadiaty RK, Herder F. 2014. MA. 2013. Breeding biology of guppy fish,
Two new species of viviparous halfbeaks Poecilia reticulata (Peters, 1895) in the
(Atherinomorhpha: Beloniformes: laboratory. Journal of the Asiatic Society of
Zenarchopteridae) endemic to Sulawesi Bangladesh Science 39: 259–267.
Tenggara, Indonesia. Raffles Bulletin of Sørensen SR, Tomkiewicz J, Munk P, Butts
Zoology 62: 200–209. IAE, Nielsen A, Lauesen P, Graver C. 2016.
Kanai T, Nanjo K, Kohno H, Sano M. 2017. Ontogeny and growth of early stages of
Ontogenetic and seasonal changes in the diet captive-bred European eel. Aquaculture 456:
of the halfbeak Zenarchopterus dunckeri at 50–61.
Iriomote Island, southern Japan. Ichthyological Tabassum S, Yousuf F, Elahi N, Hossain MY,
Research 64: 470–474. Hossen MA, Nawer F, Khatun D, Parvin MF.
Kobayashi H, Masengi WA, Yamahira K. 2020. 2017. Diet and feeding ecology of black-
A new “beakless” halfbeak of the Genus barred halfbeak Hemiramphus far (Forsskal
Nomorhamphus from Sulawesi (Teleostei: 1775) (Hemiramphidae) from Karachi coast
Zenarchopteridae). Copeia 108: 522–531. of Pakistan. Journal of Coastal Life Medicine
Kraemer J, Hadiaty RK, Herder F. 2019a. 5: 4–6.
Nomorhamphus versicolor, a new species Tinguely SM, Lange A, Tyler CR. 2019. Ontogeny
of bluntnosed halfbeak from a tributary of and dynamics of the gonadal development,
the Palu River, Sulawesi Tengah (Teleostei: embryogenesis, and gestation in Xenotoca
Zenarchopteridae). Ichthyological Exploration eiseni (Cyprinidontiformes, Goodeidae).
of Freshwaters 1105: 1–8. Sexual Development 13: 297–310.
Kraemer J, Thieme P, Hadiaty RK, Herder F. Wicaksono AW, Carman O, Soelistyowati
2019b. Structure pf the andropodium of the DT, Arfah H, Alimuddin, Herjayanto M,
viviparous halfbeak genus Nomorhamphus Solahudin EA, Ahmadi K. 2022. Biometric
(Atherinomorpha:Beloniformes:Zenarchopter characterization and morphogenesis of Anasa
idae), endemic to Sulawesi, Indonesia. Raffles ornamental fish (Nomorhamphus sp.) endemic
Bulletin of Zooloogy 67: 247–259. to Lake Lindu, Central Sulawesi, Indonesia.
Ogino Y, Katoh H, Yamada G. 2004. Androgen AACL Bioflux 15: 2479–2487.