Morphometric and genetic variations of species composers of nike fish assemblages in Gorontalo Bay Waters, Indonesia
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FEMY M. SAHAMI
♥
https://orcid.org/0000-0002-6273-3109
RENE CHARLES KEPEL
https://orcid.org/0000-0001-5003-9462
ABDUL HAFIDZ OLII
SILVESTER BENNY PRATASIK
https://orcid.org/0000-0002-3765-509X
RIDWAN LASABUDA
https://orcid.org/0000-0002-4122-8103
ADNAN WANTASEN
https://orcid.org/0000-0002-6922-8295
SITTY AINSYAH HABIBIE
https://orcid.org/0000-0001-8522-9741
Abstract
Abstract. Sahami FM, Kepel RC, Olii AH, Pratasik SB, Lasabuda R, Wantasen A, Habibie SA. 2020. Morphometric and genetic variations of species composers of nike fish assemblages in Gorontalo Bay Waters, Indonesia. Biodiversitas 21: 4571-4581. Nike is Gobioidei fish at the post-larval and juvenile stages whose habitat is still in seawater before they migrate to the freshwater, grow up, and spawn to fulfill their life cycle as an amphidromous species. This study aims to identify the species composers of Nike fish in Gorontalo Bay waters based on their morphometric and molecular characters. 2,523 samples were collected from the catches of fishermen during three periods of their appearance in Gorontalo Bay waters from January to March 2019. The samples were grouped based on their similarity of melanophore patterns, and morphometric characters of 10 units were then measured. This study found 20 different groups of melanophore patterns; 15 of them were new melanophore patterns. DNA samples from each group of new melanophore patterns were then isolated for molecular analysis. The morphometric analysis grouped the 20 melanophore patterns into three separate clusters that were confirmed through molecular analysis. The results of Gen Cytochrome Oxidase I (COI) sequences indicate that the new melanophore patterns of Nike fish assemblages consisting of six species; five species in the Gobiidae family (i.e. Sicyopterus parvei, S. cynocephalus, S. longifilis, S. lagocephalus, and Stiphodon semoni) and a species in the Eleotridae family (Belobranchus belobranchus).
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References
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Dahruddin H, Hutama A, Busson F, Sauri S, Hanner R, Keith P, Hadiaty R, Hubert N. 2016. Revisiting the ichthyodiversity of Java and Bali through DNA barcodes: taxonomic coverage, identification accuracy, cryptic diversity and identification of exotic species. Molecular Ecology Resources 17(2): 1–12.
Elliott NG, Haskard K, Koslow JA. 1995. Morphometric analysis of orange roughy (Hoplostethus atalanticus) off the continental slope of southern Australia. J. Fish Biol 46(2): 202–220.
Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial Cytochrome C Oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3(5): 294–299.
Gani A, Bakri AA, Adriany DT, Nurjirana N, Herjayanto M, Bungalim MI, Ndobe S, Burhanuddin AI. 2019. Identification of freshwater goby species from The Biak and Koyoan Rivers, Luwuk Banggai, Central Sulawesi. Jurnal Ilmu Kelautan Spermonde 5(2): 57–60.
Haryono, Tjakrawidjaja AH, Riyanto A. 2002. Ichtyofauna of Bogani Nani Wartabone National Park Waters North Sulawesi. Jurnal Iktiologi Indonesia 2(2): 31–40. [Indonesian]
Hebert PDN, Stoeckle MY, Zemlak TS, Francis CM. 2004. Identification of birds through DNA barcodes. PLoS Biol 2(10): e312.
Hubert N, Hanner R, Holm E, Mandrak NE, Taylor E, Burridge M, Watkinson D, Dumont P, Curry A, Bentzen P, Zhang J, April J, Bernatchez L. 2008. Identifying Canadian freshwater fishes through DNA barcodes. PLoS One 3(6): e2490.
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Jeon HB, Choi SH, Suk HY. 2012. Exploring the utility of partial cytochrome c oxidase subunit 1 for DNA barcoding of gobies. Anim. Syst. Evol. Divers. 28(4): 269–278.
Jin X, Wang R, Wei T, Tang D, Xu T. 2013. Complete mitochondrial genome sequence of Tridentiger bifasciatus and Tridentiger barbatus (Perciformes, Gobiidae): a mitogenomic perspective on the phylogenetic relationships of Gobiidae. Mol. Biol. Rep. 42: 253–265.
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Keith P, Galewski T, Cattaneo-Berrebi G, Hoareau T, Berrebi P. 2005. Ubiquity of Sicyopterus lagocephalus (Teleostei: Gobioidei) and phylogeography of the genus Sicyopterus in the Indo-Pacific area inferred from mitochondrial cytochrome b gene. Molecular Phylogenetics and Evolution 37: 721–732.
Keith P, Hoareau TB, Lord C, Ah-Yane O, Gimonneau G, Robinet T, Valade P. 2008. Characterisation of post-larval to juvenile stages, metamorphosis and recruitment of an amphidromous goby, Sicyopterus lagocephalus (Pallas) (Teleostei?: Gobiidae?: Sicydiinae). Marine and Freshwater Research 59: 876–889.
Keith P, Lord C, Lorion J, Watanabe S, Tsukamoto K, Couloux A, Dettai A. 2011. Phylogeny and biogeography of Sicydiinae (Teleostei: Gobiidae) inferred from mitochondrial and nuclear genes. Mar. Biol. 158: 311–326.
Keith P, Lord C, Busson F, Sauri S, Hubert N, Hadiaty RK. 2015. A new species of Sicyopterus (Gobiidae) from Indonesia. Cybium 39(4): 243–248.
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Larmuseau MHD, Vancampenhout K, Raeymaekers JAM, Houdt JKJ, Volckaert FAM. 2010. Differential modes of selection on the rhodopsin gene in coastal Baltic and North Sea populations of the sand goby, Pomatoschistus minutus. Molecular Ecology 19(11): 2256–2268.
Lord C, Bellec L, Dettaï A, Bonillo C, Keith P. 2019. Does your lip stick? Evolutionary aspects of the mouth morphology of the Indo?Pacific clinging goby of the Sicyopterus genus (Teleostei: Gobioidei: Sicydiinae) based on mitogenome phylogeny. Journal of Zoological Systematic and Evolutionary Research 2019;00: 1–16.
Maeda K, Tan HH. 2013. Review of Stiphodon (Gobiidae: Sicydiinae) from Western Sumatra, with Description of a New Species. The Raffles Bulletin of Zoology 61(2): 749–761.
Maie T, Wilson M, Schoenfuss H, Blob R. 2009. Feeding kinematics and performance of Hawaiian stream Gobies, Awaous guamensis and Lentipes concolor: linkage of functional morphology and ecology. J Morphol. 270: 344–356.
Milton DA. 2009. Living in two worlds: diadromous fishes, and factors affecting population connectivity between tropical rivers and coasts. Nagelkerken I (eds) Ecological Connectivity among Tropical Coastal Ecosystems. Springer Science and Business Media B.V.
Nurjirana, Burhanuddin AI, Haris A. 2019. Diversity of penja fish (amphidromous goby) in Leppangan River, West Sulawesi, Indonesia. AACL Bioflux 12(1): 246–249.
Nurjirana, Haris A, Sahami FM, Keith P, Burhanuddin AI. 2019. Preliminary note on the morphological characters of penja (amphidromous goby post larvae) in West Sulawesi and Gorontalo Bay. IOP Conference Series: Earth and Environmental Science 370(1): 1–7.
Olii AH, Sahami FM, Hamzah SN, Pasisingi N. 2017. Preliminary findings on distribution pattern of larvae of nike fish (Awaous sp.) in the estuary of Bone River, Gorontalo Province, Indonesia. AACL Bioflux 10: 1110–1118.
Olii AH, Sahami FM, Hamzah SN, Pasisingi N. 2019. Molecular approach to identify gobioid fishes, "Nike" and "Hundala" (local name), from Gorontalo Waters, Indonesia. Online Journal of Biological Sciences 19(1): 51–56.
Pasisingi N, Abdullah S. 2018. Pattern of nike fish (gobiidae) occurrence in the Gorontalo Bay, Indonesia. Depik, Jurnal Ilmu-Ilmu Perairan, Pesisir dan Perikanan 7: 111–118.
Pasisingi N, Habibie SA, Olii AH. 2020. Are Awaous ocellaris and Belobranchus belobranchus the two species of Nike fish schools?. Aceh Journal of Animal Science 5(2): 87–91.
Roesma DI, Tjong DH, Munir W, Aidil DR. 2018. New record species of Puntius (Pisces: Cyprinidae) from West Sumatra based on cytochrome oxidase 1 gene. International Journal on Advance Science Engineering Information Technology 8: 250–256.
Roesma DI, Tjong DH, Karlina W, Aidil DR. 2019. Taxonomy confirmation of Puntius cf. binotatus from Gunung Tujuh Lake, Jambi, Indonesia based on cytochrome oxidase-I (COI) gene. Biodiversitas 20(1): 54–60.
Roesma DI, Tjong DH, Aidil DR. 2020. Phylogenetic analysis of transparent gobies in three Sumatran lakes, inferred from mitochondrial cytochrome oxidase I (COI) gene. Biodiversitas 21(1): 43–48.
Rukmana RM, Trijoko, Pratiwi R. 2014. The diversity of order perciformes fish members in Sukamade River Meru Betiri National Park. Biomedika 7(2): 46–52. [Indonesian]
Sahami FM, Kepel RC, Olii AH, Pratasik SB. 2019. Determination of morphological alteration based on molecular analysis and melanophore pattern of the migrating Nike fish in Gorontalo Bay, Indonesia. AACL Bioflux 12(4): 1358–1365.
Sahami FM, Kepel RC, Olii AH, Pratasik SB. 2019a. What species make up the Nike fish assemblages at the macrotidal estuary in Gorontalo Bay, Indonesia? [version 1; peer review: 2 approved with reservation]. F1000Research 8: 1654.
Salam A, Sahami FM, Panigoro C. 2016. Nike (Awaous melanocephalus) fishery and mercury contamination in the estuary of Bone-Bolango River. Omni-Akuatika 12(2): 130–136.
Sara L, Muskita WH, Astuti O, Safilu (2016) The reproductive biology of blue swimming crab Portunus pelagicus in Southeast Sulawesi Waters, Indonesia. AACL Bioflux 9(5): 1101–1112.
Strüder-Kypke M, Lynn DH. 2010. Comparative analysis of the mitochondrial cytochrome c oxidase subunit I (COI) gene in ciliates (Alveolata, Ciliophora) and evaluation of its suitability as a biodiversity marker. Systematics and Biodiversity 8(1): 131–148.
Taillebois L, Castelin M, Lord C, Chabarria R, Dettaï A, Keith P. 2014. New sicydiinae phylogeny (Teleostei: Gobioidei) inferred from mitochondrial and nuclear genes: insights on systematics and ancestral areas. Molecular Phylogenetics and Evolution 70: 260–271.
Teletchea F. 2009. Molecular identification methods of fish species: reassessment and possible applications. Rev. Fish Biol. Fish 19: 265–293.
Tjakrawidjaja AH. 2002. Freshwater fishes of Manggabai Flores, with the Various Limnology Aspect. Jurnal Iktiologi Indonesia 2(2): 15–22.
Thacker CE, Roje DM. 2011. Phylogeny of Gobiidae and identification of gobiid lineages. Systematics and Biodiversity 9(4): 329–347.
Thomas Jr RC, Beldia III PD, Campos WL, Santos MD. 2013. Resolving the identity of larval fishes, Dulong, in the Verde Island Passages. Philippines. Phil. J. of Nat 18: 29–36.
Usman MY. 2016. Analysis of genetic variation penja indigenous from Polewali Mandar and Nike (Awaous sp.) indigenous from Gorontalo. [Thesis]. UIN Alauddin Makassar, Makassar. [Indonesian]
Valade P, Lord C, Grondin H, Bosc P, Taillebois L, Iida M, Tsukamoto K, Keith P. 2009. Early life history and description of larval stages of an amphidromous goby, Sicyopterus lagocephalus (Gobioidei: Sicydiinae). Cybium 33(4): 309–319.
Wang ZD, Liao J, Huang CQ, Long SS, Zhang S, Guo YS, Liu L, Liu CW. 2017. Significant genetic differentiation of Gobiopterus lacustris, a newly recorded transparent goby in China. Mitochondrial DNA Part A: 1–7.
Watanabe S, Iida M, Hagihara S, Endo H, Matsuura K, Tsukamoto K. First collection of amphidromous goby post-larvae of Sicyopterus japonicus in the ocean off Shikoku, Japan. Cybium 35(4): 371–379.
Watson RE. 2008. A new species of Stiphodon from southern Sumatra (Pisces: Gobioidei: Sicydiinae). Zootaxa 1715: 43–56.
Yamasaki N, Kondo M, Maeda K, Tachihara K. 2011. Reproductive biology of three amphidromous gobies, Sicyopterus japonicus, Awaous melanocephalus, and Stenogobius sp., on Okinawa Island. Cybium, International Journal of Ichthyology 35(4): 345–360.
Dahruddin H, Hutama A, Busson F, Sauri S, Hanner R, Keith P, Hadiaty R, Hubert N. 2016. Revisiting the ichthyodiversity of Java and Bali through DNA barcodes: taxonomic coverage, identification accuracy, cryptic diversity and identification of exotic species. Molecular Ecology Resources 17(2): 1–12.
Elliott NG, Haskard K, Koslow JA. 1995. Morphometric analysis of orange roughy (Hoplostethus atalanticus) off the continental slope of southern Australia. J. Fish Biol 46(2): 202–220.
Folmer O, Black M, Hoeh W, Lutz R, Vrijenhoek R. 1994. DNA primers for amplification of mitochondrial Cytochrome C Oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3(5): 294–299.
Gani A, Bakri AA, Adriany DT, Nurjirana N, Herjayanto M, Bungalim MI, Ndobe S, Burhanuddin AI. 2019. Identification of freshwater goby species from The Biak and Koyoan Rivers, Luwuk Banggai, Central Sulawesi. Jurnal Ilmu Kelautan Spermonde 5(2): 57–60.
Haryono, Tjakrawidjaja AH, Riyanto A. 2002. Ichtyofauna of Bogani Nani Wartabone National Park Waters North Sulawesi. Jurnal Iktiologi Indonesia 2(2): 31–40. [Indonesian]
Hebert PDN, Stoeckle MY, Zemlak TS, Francis CM. 2004. Identification of birds through DNA barcodes. PLoS Biol 2(10): e312.
Hubert N, Hanner R, Holm E, Mandrak NE, Taylor E, Burridge M, Watkinson D, Dumont P, Curry A, Bentzen P, Zhang J, April J, Bernatchez L. 2008. Identifying Canadian freshwater fishes through DNA barcodes. PLoS One 3(6): e2490.
Hubert N, Kadarusman, Wibowo A, Busson F, Caruso D, Sulandari S, Nafiqoh N, Poiyaud L, Ruber L, Avarre JC, Herder F, Hanner R, Keith P, Hadiaty RK (2015) DNA Barcoding Indonesian freshwater fishes: challenges and prospects. DNA Barcodes 3: 144–169.
Ivanova NV, Zemlak TS, Hanner RH, Hebert PDN. 2007. Universal primer cocktails for fish DNA barcoding. Molecular Ecology Notes 7(4): 544–548.
Jeon HB, Choi SH, Suk HY. 2012. Exploring the utility of partial cytochrome c oxidase subunit 1 for DNA barcoding of gobies. Anim. Syst. Evol. Divers. 28(4): 269–278.
Jin X, Wang R, Wei T, Tang D, Xu T. 2013. Complete mitochondrial genome sequence of Tridentiger bifasciatus and Tridentiger barbatus (Perciformes, Gobiidae): a mitogenomic perspective on the phylogenetic relationships of Gobiidae. Mol. Biol. Rep. 42: 253–265.
Keith P. 2003. Biology and ecology of amphidromous Gobiidae of the Indo-Pacific and the Caribbean regions. Journal of Fish Biology 63: 831–847.
Keith P, Galewski T, Cattaneo-Berrebi G, Hoareau T, Berrebi P. 2005. Ubiquity of Sicyopterus lagocephalus (Teleostei: Gobioidei) and phylogeography of the genus Sicyopterus in the Indo-Pacific area inferred from mitochondrial cytochrome b gene. Molecular Phylogenetics and Evolution 37: 721–732.
Keith P, Hoareau TB, Lord C, Ah-Yane O, Gimonneau G, Robinet T, Valade P. 2008. Characterisation of post-larval to juvenile stages, metamorphosis and recruitment of an amphidromous goby, Sicyopterus lagocephalus (Pallas) (Teleostei?: Gobiidae?: Sicydiinae). Marine and Freshwater Research 59: 876–889.
Keith P, Lord C, Lorion J, Watanabe S, Tsukamoto K, Couloux A, Dettai A. 2011. Phylogeny and biogeography of Sicydiinae (Teleostei: Gobiidae) inferred from mitochondrial and nuclear genes. Mar. Biol. 158: 311–326.
Keith P, Lord C, Busson F, Sauri S, Hubert N, Hadiaty RK. 2015. A new species of Sicyopterus (Gobiidae) from Indonesia. Cybium 39(4): 243–248.
Landau S, Everit BS. 2004. A Handbook of Statistical Analyses using SPSS. Chapman & Hall/CRC, Washington DC.
Larmuseau MHD, Vancampenhout K, Raeymaekers JAM, Houdt JKJ, Volckaert FAM. 2010. Differential modes of selection on the rhodopsin gene in coastal Baltic and North Sea populations of the sand goby, Pomatoschistus minutus. Molecular Ecology 19(11): 2256–2268.
Lord C, Bellec L, Dettaï A, Bonillo C, Keith P. 2019. Does your lip stick? Evolutionary aspects of the mouth morphology of the Indo?Pacific clinging goby of the Sicyopterus genus (Teleostei: Gobioidei: Sicydiinae) based on mitogenome phylogeny. Journal of Zoological Systematic and Evolutionary Research 2019;00: 1–16.
Maeda K, Tan HH. 2013. Review of Stiphodon (Gobiidae: Sicydiinae) from Western Sumatra, with Description of a New Species. The Raffles Bulletin of Zoology 61(2): 749–761.
Maie T, Wilson M, Schoenfuss H, Blob R. 2009. Feeding kinematics and performance of Hawaiian stream Gobies, Awaous guamensis and Lentipes concolor: linkage of functional morphology and ecology. J Morphol. 270: 344–356.
Milton DA. 2009. Living in two worlds: diadromous fishes, and factors affecting population connectivity between tropical rivers and coasts. Nagelkerken I (eds) Ecological Connectivity among Tropical Coastal Ecosystems. Springer Science and Business Media B.V.
Nurjirana, Burhanuddin AI, Haris A. 2019. Diversity of penja fish (amphidromous goby) in Leppangan River, West Sulawesi, Indonesia. AACL Bioflux 12(1): 246–249.
Nurjirana, Haris A, Sahami FM, Keith P, Burhanuddin AI. 2019. Preliminary note on the morphological characters of penja (amphidromous goby post larvae) in West Sulawesi and Gorontalo Bay. IOP Conference Series: Earth and Environmental Science 370(1): 1–7.
Olii AH, Sahami FM, Hamzah SN, Pasisingi N. 2017. Preliminary findings on distribution pattern of larvae of nike fish (Awaous sp.) in the estuary of Bone River, Gorontalo Province, Indonesia. AACL Bioflux 10: 1110–1118.
Olii AH, Sahami FM, Hamzah SN, Pasisingi N. 2019. Molecular approach to identify gobioid fishes, "Nike" and "Hundala" (local name), from Gorontalo Waters, Indonesia. Online Journal of Biological Sciences 19(1): 51–56.
Pasisingi N, Abdullah S. 2018. Pattern of nike fish (gobiidae) occurrence in the Gorontalo Bay, Indonesia. Depik, Jurnal Ilmu-Ilmu Perairan, Pesisir dan Perikanan 7: 111–118.
Pasisingi N, Habibie SA, Olii AH. 2020. Are Awaous ocellaris and Belobranchus belobranchus the two species of Nike fish schools?. Aceh Journal of Animal Science 5(2): 87–91.
Roesma DI, Tjong DH, Munir W, Aidil DR. 2018. New record species of Puntius (Pisces: Cyprinidae) from West Sumatra based on cytochrome oxidase 1 gene. International Journal on Advance Science Engineering Information Technology 8: 250–256.
Roesma DI, Tjong DH, Karlina W, Aidil DR. 2019. Taxonomy confirmation of Puntius cf. binotatus from Gunung Tujuh Lake, Jambi, Indonesia based on cytochrome oxidase-I (COI) gene. Biodiversitas 20(1): 54–60.
Roesma DI, Tjong DH, Aidil DR. 2020. Phylogenetic analysis of transparent gobies in three Sumatran lakes, inferred from mitochondrial cytochrome oxidase I (COI) gene. Biodiversitas 21(1): 43–48.
Rukmana RM, Trijoko, Pratiwi R. 2014. The diversity of order perciformes fish members in Sukamade River Meru Betiri National Park. Biomedika 7(2): 46–52. [Indonesian]
Sahami FM, Kepel RC, Olii AH, Pratasik SB. 2019. Determination of morphological alteration based on molecular analysis and melanophore pattern of the migrating Nike fish in Gorontalo Bay, Indonesia. AACL Bioflux 12(4): 1358–1365.
Sahami FM, Kepel RC, Olii AH, Pratasik SB. 2019a. What species make up the Nike fish assemblages at the macrotidal estuary in Gorontalo Bay, Indonesia? [version 1; peer review: 2 approved with reservation]. F1000Research 8: 1654.
Salam A, Sahami FM, Panigoro C. 2016. Nike (Awaous melanocephalus) fishery and mercury contamination in the estuary of Bone-Bolango River. Omni-Akuatika 12(2): 130–136.
Sara L, Muskita WH, Astuti O, Safilu (2016) The reproductive biology of blue swimming crab Portunus pelagicus in Southeast Sulawesi Waters, Indonesia. AACL Bioflux 9(5): 1101–1112.
Strüder-Kypke M, Lynn DH. 2010. Comparative analysis of the mitochondrial cytochrome c oxidase subunit I (COI) gene in ciliates (Alveolata, Ciliophora) and evaluation of its suitability as a biodiversity marker. Systematics and Biodiversity 8(1): 131–148.
Taillebois L, Castelin M, Lord C, Chabarria R, Dettaï A, Keith P. 2014. New sicydiinae phylogeny (Teleostei: Gobioidei) inferred from mitochondrial and nuclear genes: insights on systematics and ancestral areas. Molecular Phylogenetics and Evolution 70: 260–271.
Teletchea F. 2009. Molecular identification methods of fish species: reassessment and possible applications. Rev. Fish Biol. Fish 19: 265–293.
Tjakrawidjaja AH. 2002. Freshwater fishes of Manggabai Flores, with the Various Limnology Aspect. Jurnal Iktiologi Indonesia 2(2): 15–22.
Thacker CE, Roje DM. 2011. Phylogeny of Gobiidae and identification of gobiid lineages. Systematics and Biodiversity 9(4): 329–347.
Thomas Jr RC, Beldia III PD, Campos WL, Santos MD. 2013. Resolving the identity of larval fishes, Dulong, in the Verde Island Passages. Philippines. Phil. J. of Nat 18: 29–36.
Usman MY. 2016. Analysis of genetic variation penja indigenous from Polewali Mandar and Nike (Awaous sp.) indigenous from Gorontalo. [Thesis]. UIN Alauddin Makassar, Makassar. [Indonesian]
Valade P, Lord C, Grondin H, Bosc P, Taillebois L, Iida M, Tsukamoto K, Keith P. 2009. Early life history and description of larval stages of an amphidromous goby, Sicyopterus lagocephalus (Gobioidei: Sicydiinae). Cybium 33(4): 309–319.
Wang ZD, Liao J, Huang CQ, Long SS, Zhang S, Guo YS, Liu L, Liu CW. 2017. Significant genetic differentiation of Gobiopterus lacustris, a newly recorded transparent goby in China. Mitochondrial DNA Part A: 1–7.
Watanabe S, Iida M, Hagihara S, Endo H, Matsuura K, Tsukamoto K. First collection of amphidromous goby post-larvae of Sicyopterus japonicus in the ocean off Shikoku, Japan. Cybium 35(4): 371–379.
Watson RE. 2008. A new species of Stiphodon from southern Sumatra (Pisces: Gobioidei: Sicydiinae). Zootaxa 1715: 43–56.
Yamasaki N, Kondo M, Maeda K, Tachihara K. 2011. Reproductive biology of three amphidromous gobies, Sicyopterus japonicus, Awaous melanocephalus, and Stenogobius sp., on Okinawa Island. Cybium, International Journal of Ichthyology 35(4): 345–360.
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