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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Superclassis/Classis: Actinopterygii
Classis/Subclassis: Actinopteri
Subclassis/Infraclassis: Neopterygii
Infraclassis: Teleostei
Megacohors: Osteoglossocephalai
Supercohors: Clupeocephala
Cohors: Euteleosteomorpha
Subcohors: Neoteleostei
Infracohors: Eurypterygia
Sectio: Ctenosquamata
Subsectio: Acanthomorphata
Divisio/Superordo: Acanthopterygii
Subdivisio: Percomorphaceae
Series: Eupercaria
Ordo: Perciformes
Subordo: Percoidei
Superfamilia: Percoidea

Familia: Sciaenidae
Genus: Johnius
Species (33): J. amblycephalus – J. australis – J. belangerii – J. borneensis – J. cantori – J. carouna – J. carutta – J. coitor – J. distinctus – J. dorsalis – J. dussumieri – J. elongatus – J. fasciatus – J. fuscolineatus – J. gangeticus – J. glaucus – J. goldmani – J. grypotus – J. heterolepis – J. hypostoma – J. laevis – J. latifrons – J. macropterus – J. macrorhynus – J. majan – J. mannarensis – J. novaeguineae – J. novaehollandiae – J. pacificus – J. philippinus – J. plagiostoma – J. sina – J. trachycephalus – J. trewavasae
Name

Johnius Bloch, 1793
References

Bloch, M.E. 1793. Naturgeschichte der ausländischen Fische. Berlin. v. 7: i–xiv + 1–144, Pls. 325–360. Reference page.
Chao, N.L., Chang, C-W., Chen, M-H, Guo, C-C., Lin, B-A., Liou, Y-Y., Shen, K-N. & Liu, M. 2019. Johnius taiwanensis, a new species of Sciaenidae from the Taiwan Strait, with a key to Johnius species from Chinese waters. Zootaxa 4651(2): 259–270. DOI: 10.11646/zootaxa.4651.2.3 Paywall Reference page.
Zhao, L-L., Yi, D., Li, C-H., Sun, D-R., Xu, H-X. & Gao, T-X. 2017. Phylogeography and population structure of - grypotus (Richardson, 1846) as revealed by mitochondrial control region sequences. ZooKeys 705: 143—158. DOI: 10.3897/zookeys.705.13001. Reference page.

Johnius is a genus of marine ray-finned fishes belonging to the family Sciaenidae, the drums and croakers. They are commonly known as croakers due to their ability to produce purring, croaking and knocking sounds. The sounds are produced mainly at night and are thought to be either involved in defense or for courtship.[2]
Taxonomy

The genus name was erected by Marcus Bloch in 1793 based on a specimen obtained from Tranquebar from Reverend Christoph Samuel John which was named as Johnius carutta. There are about 36 species in the genus, all within the Indo-West Pacific waters.[3] This genus has been placed in the subfamily Otolithinae by some workers,[4] but the 5th edition of Fishes of the World does not recognise subfamilies within the Sciaenidae which it places in the order Acanthuriformes.[5]

Two subgenera are recognised within the genus Johnius, the nominate subgenus has an inferior mouth with the teeth on the lower jaw being uniform in size, although there may be a small number of molar-like teeth towards the rear of the jaw. They also have the teeth on the rear of the upper jaw relatively closely set. The subgenus Johnieops has a terminal or subterminal mouth with an inner row of enlarged teeth on the lower jaw and the outer teeth on the upper jaw are relatively widely spaced.[6] The type species of Johnieops is Sciaena osseus Day, 1876.[1] However, the two subgenera may not represent monophyletic groupings.[6]
Etymology

Johnius honours the Danish missionary Christoph Samuel John who collected specimens at Tranquebar for Bloch, presumed to include the types of Johnius carutta and Pennahia aneus.[7]
Species

Johnius includes 36 species,[8] divided into two subgenera:[7]

Subgenus Johnieops
Johnius borneensis (Bleeker, 1851) (Sharpnose hammer croaker)
Johnius distinctus (Tanaka, 1916)
Johnius dorsalis (Peters, 1855) (Small kob)
Johnius dussumieri (Cuvier, 1830) (Sin croaker)
Johnius pacificus Hardenberg, 1941 (Pacific croaker)
Johnius philippinus Sasaki, 1999
Johnius plagiostoma (Bleeker, 1849) (Large-eye croaker)
Subgenus Johnius
Johnius amblycephalus (Bleeker, 1855) (Bearded croaker)
Johnius australis (Günther, 1880) (Bottlenose jewfish)
Johnius belangerii (Cuvier, 1830) (Belanger's croaker)
Johnius cantori Bleeker, 1874
Johnius carouna (Cuvier, 1830) (Caroun croaker)
Johnius carutta Bloch, 1793 (Karut croaker)
Johnius coitor (Hamilton, 1822) (Coitor croaker)
Johnius elongatus Lal Mohan, 1976 (Spindle croaker)
Johnius fasciatus Chu, Lo & Wu, 1963
Johnius fuscolineatus (von Bonde, 1923) (Bellfish)
Johnius gangeticus Talwar, 1991
Johnius glaucus (Day, 1876) (Pale spotfin croaker)
Johnius goldmani (Bleeker, 1855)
Johnius grypotus (Richardson, 1846)
Johnius heterolepis Bleeker, 1873 (Large-scale croaker)
Johnius hypostoma (Bleeker, 1854) (Small-mouth croaker)
Johnius laevis Sasaki & Kailola, 1991 (Smooth croaker)
Johnius latifrons Sasaki, 1992 (Broad-head croaker)
Johnius macropterus (Bleeker, 1853) (Largefin croaker)
Johnius macrorhynus (Lal Mohan, 1976) (Big-snout croaker)
Johnius majan Iwatsuki, Jawad & Al-Mamry, 2012 (Majan croaker)
Johnius mannarensis Lal Mohan, 1971 (Mannar croaker)
Johnius novaeguineae (Nichols, 1950) (Paperhead croaker)
Johnius novaehollandiae (Steindachner, 1866)
Johnius sasakii Hanafi, Chen, Seah, Chang, Liu & Chao, 2022[9]
Johnius taiwanensis Chao, Chang, Chen, Guo, Lin, Liou, Shen & Liu, 2019
Johnius trachycephalus (Bleeker, 1851) (Leaftail croaker)
Johnius trewavasae Sasaki, 1992 (Trewavas croaker)
Johnius weberi Hardenberg, 1936 (Weber's croaker)

Some authorities recognise Johnius sina as a valid species,[10] while others treat it as a synonym of J. dussumieri.[8]
Characteristics

Johnius croakers are relatively small Scaienids, typically less than 30 cm (12 in). They have a characteristic hammer-shaped swim bladder with between 12 and 20 pairs of dendritic appendages along its sides.[6] The first lateral appendage extends to the dorsal corner of the gill opening. They have large paired sagittal otoliths that are triangular. These fishes typically have a small mouth that is positioned sub-terminally to inferiorly. A few species have a short barbel on the chin but most species have no barbel on the chin.[3] The largest species as J. dorsalis and J. dussumieri which both have a maximum published total length of 40 cm (16 in).[8]
Distribution

Johnius croakers are found in the Indo-Pacific region from eastern Africa to Australia and New Guinea.[8]
References

Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Genera in the family Sciaenidae". Catalog of Fishes. California Academy of Sciences. Retrieved 12 May 2023.
Lin, Yuan C.; Mok, Hin K.; Huang, Bao Q. (2007). "Sound characteristics of big-snout croaker, Johnius macrorhynus (Sciaenidae)". The Journal of the Acoustical Society of America. 121 (1): 586–593. doi:10.1121/1.2384844. ISSN 0001-4966. PMID 17297812.
Chao, Ning Labbish; Chang, Chih-Wei; Chen, Meng-Hsien; Guo, Chang-Chang; Lin, Bai-An; Liou, You-Yu; Shen, Kang-Ning; Liu, Min (2019). "Johnius taiwanensis, a new species of Sciaenidae from the Taiwan Strait, with a key to Johnius species from Chinese waters". Zootaxa. 4651 (2): 259–270. doi:10.11646/zootaxa.4651.2.3. ISSN 1175-5334. S2CID 201210739.
Kunio Sasaki (1989). "Phylogeny of the family Sciaenidae, with notes on its Zoogeography (Teleostei, Peciformes)" (PDF). Memoirs of the Faculty of Fishes Hokkaido University. 36 (1–2): 1–137.
J. S. Nelson; T. C. Grande; M. V. H. Wilson (2016). Fishes of the World (5th ed.). Wiley. pp. 497–502. ISBN 978-1-118-34233-6.
Kunio Sasaki (2022). "Family Sciaenidae Croakers, Drums and Cobs". In Phillip C Heemstra; Elaine Heemstra; David A Ebert; Wouter Holleman; John E Randall (eds.). Coastal Fishes of the Western Indian Ocean Volume 3 (PDF). South African Institute for Aquatic Biodiversity. pp. 389–414. ISBN 978-1-990951-30-5.
Christopher Scharpf & Kenneth J. Lazara, eds. (12 January 2024). "Order ACANTHURIFORMES (part 5): Family SCIAENIDAE". The ETYFish Project Fish Name Etymology Database. v. 6.0. Christopher Scharpf and Kenneth J. Lazara. Retrieved 24 March 2024.
Froese, Rainer and Pauly, Daniel, eds. (2023). Species of Johnius in FishBase. February 2023 version.
Norhafiz Hanafi; Meng-Hsien Chen; Yin-Giat Seah; et al. (2022). "Johnius sasakii, a new species of croaker (Perciformes: Sciaenidae) with a key to Johnius from East Malaysia, Borneo". Zootaxa. 5116 (3): 393–409. doi:10.11646/ZOOTAXA.5116.3.5. PMID 35391328. S2CID 247591447.
Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Species in the genus Johnius". Catalog of Fishes. California Academy of Sciences. Retrieved 16 May 2023.

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