Chaetodon Melannotus

''Chaetodon'' is a tropical fish genus in the family Chaetodontidae. Like their relatives, they are known as "butterflyfish". This genus is by far the largest among the Chaetodontidae, with about 90 living species included here, though most might warrant recognition as distinct genera.

Species

Extant species

There are currently 87 recognized species in this genus:

''Chaetodon sensu stricto''

* '' Chaetodon capistratus'' Linnaeus, 1758 (Foureye butterflyfish)
* ''Chaetodon ocellatus'' Bloch, 1787 (Spotfin butterflyfish)
* '' Chaetodon striatus'' Linnaeus, 1758 (Banded butterflyfish)

''C. robustus'' group

* '' Chaetodon hoefleri'' Steindachner, 1881 (Four-banded butterflyfish)
* '' Chaetodon robustus'' Günther, 1860 (Three-banded butterflyfish)

''Lepidochaetodon'' group

Image:Bep chaetodon punctatofasciatus.jpg, Spotband butterflyfish
''Chaetodon (Exornator) punctatofasciatus''
Image:Chaetodon guttatissimus_01.jpg, Peppered butterflyfish
''Chaetodon (Exornator) guttatissimus''
Image:8070 aquaimages.jpg, Fourspot butterflyfish
''Chaetodon (Exornator) quadrimaculatus''
Image:Chaetodon unimaculatus.jpg, Teardrop butterflyfish
''Chaetodon (Lepidochaetodon) unimaculatus''
Image:Bep chaetodon madagaskariensis.jpg, Seychelles butterflyfish
''Chaetodon (Rhombochaetodon) madagaskariensis''
Image:Chaetodon blackburnii Réunion.jpg, Brownburnie
''Chaetodon (Rhombochaetodon) blackburnii''
Image:Chaetodon interruptus, groupe.jpg, A group of Yellow teardrop butterflyfish
''Chaetodon (Lepidochaetodon) interruptus''

Subgenus ''Exornator'' (including ''Burgessius, Heterochaetodon'' and possibly ''Rhombochaetodon''; tentatively placed here)
* '' Chaetodon citrinellus'' G. Cuvier, 1831 (Speckled butterflyfish, Citron butterflyfish)
* '' Chaetodon dolosus'' C. G. E. Ahl, 1923 (African butterflyfish)
* '' Chaetodon guentheri'' C. G. E. Ahl, 1923 (Crochet butterflyfish)
* '' Chaetodon guttatissimus'' E. T. Bennett, 1833 (Peppered butterflyfish)
* '' Chaetodon miliaris'' Quoy & Gaimard, 1825 (Millet butterflyfish)
* '' Chaetodon multicinctus'' A. Garrett, 1863 (Pebbled butterflyfish)
* '' Chaetodon pelewensis'' Kner, 1868 (Sunset butterflyfish)
* '' Chaetodon punctatofasciatus'' G. Cuvier, 1831 (Spotband butterflyfish)
* '' Chaetodon quadrimaculatus'' J. E. Gray, 1831 (Fourspot butterflyfish)
* '' Chaetodon sedentarius'' Poey, 1860 (Reef butterflyfish)

Subgenus ''Lepidochaetodon'' (including ''Tifia'')
* '' Chaetodon daedalma'' D. S. Jordan & Fowler, 1902 (Wrought-iron butterflyfish) (tentatively placed here)
* '' Chaetodon interruptus'' C. G. E. Ahl, 1923 (Yellow teardrop butterflyfish)
* '' Chaetodon kleinii'' Bloch, 1790 (Sunburst butterflyfish, black-lipped butterflyfish, Klein's butterflyfish)
* ''Chaetodon litus'' J. E. Randall & D. K. Caldwell, 1973 (Easter Island butterflyfish) (tentatively placed here)
* '' Chaetodon nippon'' Steindachner & Döderlein ( de), 1883 (Japanese butterflyfish) (tentatively placed here)
* '' Chaetodon smithi'' J. E. Randall, 1975 (Smith's butterflyfish) (tentatively placed here)
* '' Chaetodon trichrous'' Günther, 1874 (Tahiti butterflyfish)
* '' Chaetodon unimaculatus'' Bloch, 1787 (Teardrop butterflyfish)

Subgenus ''Rhombochaetodon'' (including ''Roaops'', might belong in ''Exornator''; tentatively placed here)
* '' Chaetodon argentatus'' H. M. Smith & Radcliffe, 1911 (Asian butterflyfish)
* '' Chaetodon blackburnii'' Desjardins, 1836 (Brownburnie) (tentatively placed here)
* '' Chaetodon burgessi'' G. R. Allen & Starck, 1973 (Burgess' butterflyfish)
* '' Chaetodon declivis'' J. E. Randall, 1975 (Marquesas butterflyfish) (tentatively placed here)
* '' Chaetodon flavocoronatus'' R. F. Myers, 1980 (Yellow-crowned butterflyfish) (tentatively placed here)
* '' Chaetodon fremblii'' E. T. Bennett, 1828 (Blue-striped butterflyfish)
* '' Chaetodon madagaskariensis'' C. G. E. Ahl, 1923 (Seychelles butterflyfish)
* '' Chaetodon mertensii'' G. Cuvier, 1831 (Atoll butterflyfish)
* '' Chaetodon mitratus'' Günther, 1860 (Indian butterflyfish) (tentatively placed here)
* ''Chaetodon paucifasciatus'' C. G. E. Ahl, 1923 (Eritrean butterflyfish, Crown butterflyfish)
* '' Chaetodon tinkeri'' L. P. Schultz, 1951 (Hawaiian butterflyfish) (tentatively placed here)
* '' Chaetodon xanthurus'' Bleeker, 1857 (Pearlscale butterflyfish, Philippines chevron butterflyfish, yellow-tailed butterflyfish)

''Megaprotodon'' group

Image:Chaetodon ornatissimus by NPS.jpg, Ornate butterflyfish
''Chaetodon ("Citharoedus") ornatissimus''
Image:Chaetodon reticulatus by NPS.jpg, Mailed butterflyfish
''Chaetodon ("Citharoedus") reticulatus''
Image:Bep chaetodon austriacus edit.jpg, Blacktail butterflyfish
''Chaetodon (Corallochaetodon) austriacus''
Image:Bep chaetodon lunulatus.jpg, Oval butterflyfish
''Chaetodon (Corallochaetodon) lunulatus''
Image:Bep chaetodon octofasciatus.jpg, Eightband butterflyfish
''Chaetodon (Discochaetodon) octofasciatus''
Image:Bep chaetodon baronessa.jpg, Eastern triangular butterflyfish
''Chaetodon (Gonochaetodon) baronessa''
Image:Bep chaetodon trifascialis.jpg, Chevron butterflyfish
''Chaetodon (Megaprotodon) trifascialis''
Image:Bep chaetodon speculum.jpg, Mirror butterflyfish
''Chaetodon (Tetrachaetodon) speculum''
Image:Chaetodon zanzibarensis Réunion.JPG, Zanzibar butterflyfish
''Chaetodon (Tetrachaetodon) zanzibarensis''

Subgenus ''"Citharoedus"'' (tentatively placed here)
* '' Chaetodon meyeri'' Bloch & J. G. Schneider, 1801 (Scrawled butterflyfish)
* '' Chaetodon ornatissimus'' G. Cuvier, 1831 (Ornate butterflyfish)
* '' Chaetodon reticulatus'' G. Cuvier, 1831 (Mailed butterflyfish)

Subgenus ''Corallochaetodon'' (tentatively placed here)
* '' Chaetodon austriacus'' Rüppell, 1836 (Blacktail butterflyfish, exquisite butterflyfish)
* '' Chaetodon lunulatus'' Quoy & Gaimard, 1825 (Oval butterflyfish, red-finned butterflyfish)
* '' Chaetodon melapterus'' Guichenot, 1863 (Arabian butterflyfish) (tentatively placed here)
* '' Chaetodon trifasciatus'' M. Park, 1797 (Melon butterflyfish)

Subgenus ''Discochaetodon''
* '' Chaetodon aureofasciatus'' W. J. Macleay, 1878 (Golden butterflyfish)
* '' Chaetodon octofasciatus'' Bloch, 1787 (Eightband butterflyfish)
* '' Chaetodon rainfordi'' McCulloch, 1923 (Rainford's butterflyfish)
* '' Chaetodon tricinctus'' Waite, 1901 (Three-striped butterflyfish) (tentatively placed here)

Subgenus ''Gonochaetodon'' (tentatively placed here)
* '' Chaetodon baronessa'' G. Cuvier, 1829 (Eastern triangle butterflyfish)
* '' Chaetodon larvatus'' G. Cuvier, 1831 (Hooded butterflyfish)
* '' Chaetodon triangulum'' G. Cuvier, 1831 (Triangle butterflyfish)

Subgenus ''Megaprotodon''
* '' Chaetodon trifascialis'' Quoy & Gaimard, 1825 (Chevron Butterflyfish, Triangulate butterflyfish, V-lined butterflyfish)

Subgenus ''Tetrachaetodon''
* '' Chaetodon andamanensis'' Kuiter & Debelius ( de), 1999 (Andaman butterflyfish) (tentatively placed here)
* '' Chaetodon bennetti'' G. Cuvier, 1831 (Bluelashed butterflyfish)
* '' Chaetodon plebeius'' G. Cuvier, 1831 (Blue-blotched butterflyfish)
* '' Chaetodon speculum'' G. Cuvier, 1831 (Mirror butterflyfish)
* '' Chaetodon zanzibarensis'' Playfair ( fr), 1867 (Zanzibar butterflyfish)

''Rabdophorus'' group

Including ''Aspilurochaetodon, Chaetodontops''

Image:Chaetodon auriga by NPS.jpg, Threadfin butterflyfish
''Chaetodon (Rabdophorus) auriga''
Image:Bep chaetodon falcula.jpg, Blackwedged butterflyfish
''Chaetodon (Rabdophorus) falcula''
File:Chaetodon melannotus edit4.jpg, Blackback butterflyfish
''Chaetodon (Rabdophorus) melannotus''
Image:Chaetodon lineolatus.jpg, Lined butterflyfish
''Chaetodon (Rabdophorus) lineolatus''
Image:Chaetodon collare 1.jpg, Redtail butterflyfish
''Chaetodon (Rabdophorus) collare''
Image:Chaetodon flavirostris 1.jpg, Black butterflyfish
''Chaetodon (Rabdophorus) flavirostris''
Image:Chaetodon semilarvatus by Konstanze Löwe.JPG, Bluecheek butterflyfish
''Chaetodon (Rabdophorus) semilarvatus''
Image:Chaetodon ephippium PLW edit.jpg, Saddle butterflyfish
''Chaetodon (Rabdophorus) ephippium''

* '' Chaetodon adiergastos'' Seale, 1910 (Philippine butterflyfish)
* '' Chaetodon auriga'' Forsskål, 1775 (Threadfin butterflyfish)
* '' Chaetodon auripes'' D. S. Jordan & Snyder, 1901 (Oriental butterflyfish) (tentatively placed here)
* '' Chaetodon collare'' Bloch, 1787 (Redtail butterflyfish, Pakistani butterflyfish)
* '' Chaetodon decussatus'' G. Cuvier, 1829 (Indian vagabond butterflyfish)
* '' Chaetodon ephippium'' G. Cuvier, 1831 (Saddle butterflyfish)
* '' Chaetodon falcula'' Bloch, 1795 (Blackwedged butterflyfish)
* '' Chaetodon fasciatus'' Forsskål, 1775 (Diagonal butterflyfish, Red Sea raccoon butterflyfish)
* '' Chaetodon flavirostris'' Günther, 1874 (Black butterflyfish)
* '' Chaetodon gardineri'' Norman, 1939 (Gardner's butterflyfish) (tentatively placed here)
* '' Chaetodon leucopleura'' Playfair, 1867 (Somali butterflyfish) (tentatively placed here)
* '' Chaetodon lineolatus'' G. Cuvier, 1831 (Lined butterflyfish)
* '' Chaetodon lunula'' ( Lacépède, 1802) (Raccoon butterflyfish, Crescent-masked butterflyfish, lunule butterflyfish)
* '' Chaetodon melannotus'' Bloch & J. G. Schneider, 1801 (Blackback butterflyfish)
* ''Chaetodon mesoleucos'' Forsskål, 1775 (White-faced butterflyfish)
* '' Chaetodon nigropunctatus'' Sauvage, 1880 (Black-spotted butterflyfish) (tentatively placed here)
* '' Chaetodon ocellicaudus'' G. Cuvier, 1831 (Spot-tail butterflyfish)
* '' Chaetodon oxycephalus'' Bleeker, 1853 (Spot-naped butterflyfish)
* '' Chaetodon pictus'' Forsskål, 1775 (Horseshoe butterflyfish)
* '' Chaetodon rafflesii'' Anonymous ennett 1830 (Latticed butterflyfish)
* '' Chaetodon selene'' Bleeker, 1853 (Yellow-dotted butterflyfish)
* '' Chaetodon semeion'' Bleeker, 1855 (Dotted butterflyfish)
* '' Chaetodon semilarvatus'' G. Cuvier, 1831 (Bluecheek butterflyfish)
* '' Chaetodon ulietensis'' G. Cuvier, 1831 (Pacific double-saddle butterflyfish, false falcula butterflyfish)
* '' Chaetodon vagabundus'' Linnaeus, 1758 (Vagabond butterflyfish)
* '' Chaetodon wiebeli'' Kaup, 1863 (Hong Kong butterflyfish)
* '' Chaetodon xanthocephalus'' E. T. Bennett, 1833 (Yellowhead butterflyfish) (tentatively placed here)

''Incertae sedis''

Most are probably either ''Chaetodon sensu stricto'' or ''C. robustus'' group.
* '' Chaetodon assarius'' Waite, 1905 (West Australian butterflyfish)
* '' Chaetodon dialeucos'' Salm & Mee, 1989 (Oman butterflyfish)
* '' Chaetodon humeralis'' Günther, 1860
* '' Chaetodon marleyi'' Regan, 1921 (Doublesash butterflyfish)
* '' Chaetodon sanctaehelenae'' Günther, 1868 (Saint Helena butterflyfish)

Fossil species

The following fossil species are known:

* Subgenus †''Arambourgchaetodon''
** †'' Chaetodon ficheuri'' Arambourg, 1927 (Late Miocene of Algeria)
* Subgenus †''Blumchaetodon''
** †''Chaetodon wattsi'' Marramà, Giusberti & Carnevale, 2022 (Early Oligocene of Italy)

The species '' Chaetodon penniger'' Bogachev, 1964 from the late Oligocene of Azerbaijan cannot be confidently assigned to this genus due to the incompleteness of the fossil.

Proposed subgenera

Several subgenera have been proposed for splitting out of this group. It is becoming clear how the genus might be split up, with a range of DNA sequence data in large parts agreeing with S.D. Blum's landmark 1988 phylogenetic assessment of osteology.

Basically, a core group around the type species '' Chaetodon capistratus'' would remain in ''Chaetodon'', while maybe four clades would be split off. These could use the names ''Lepidochaetodon'', ''Megaprotodon'' and ''Rabdophorus'', and there is one unnamed group containing the three-banded butterflyfish (''C. robustus'') and its relatives. But the monophyly of the ''Lepidochaetodon'' group is not fully established; it is both unclear whether ''Rhombochaetodon'' is a lineage distinct from ''Exornator'', and whether ''Lepidochaetodon'' is indeed closer to these than to any other ''Chaetodon'', particularly to some lineages otherwise placed in ''Megaprotodon''.

'' Prognathodes'', for some time contained in ''Chaetodon'', is worthy of recognition as full genus, as is ''Roa''.

Historically, more distantly related fish were placed in ''Chaetodon'' too, for resembling them in details – e.g. the common scat ''Scatophagus argus'', which has a similar shape and size, as well as armored larvae like the Chaetodontidae – or simply because they are colorful, smallish, and unusual-looking – e.g. the quite unrelated paradise fish, as ''C. chinensis''.

The classification proposed in Fessler and Westneat is based on the species of ''Chaetodon'' they sampled and these sorted into the following clades:

*Clade 1
**''Chaetodon robustus''
**''Chaetodon hoefleri''
*Clade 2
**''Chaetodon kleini''
**''Chaetodon unimaculatus''
**''Chaetodon fremblii''
**''Chaetodon burgessi''
**''Chaetodon xanthurus''
**''Chaetodon paucifasciatus''
**''Chaetodon mertensi''
**''Chaetodon quadrimaculatus''
**''Chaetodon citrinellus''
**''Chaetodon multicinctus''
**''Chaetodon punctatofasciatus''
**''Chaetodon pelewensis''
**''Chaetodon sedentarius''
**''Chaetodon dolosus''
**''Chaetodon miliaris''
**''Chaetodon guentheri''
*Clade 3
**''Chaetodon austriacus''
**''Chaetodon trifasciatus''
**''Chaetodon ornatissimus''
**''Chaetodon reticularis''
**''Chaetodon meyeri''
**''Chaetodon larvatus''
**''Chaetodon baronessa''
**''Chaetodon trifascialis''
**''Chaetodon rainfordi''
**''Chaetodon octofasciatus''
**''Chaetodon bennetti''
**''Chaetodon plebius''
**''Chaetodon speculum''
**''Chaetodon zanzibarensis''
*Clade 4
**''Chaetodon ocellatus''
**''Chaetodon capistratus''
**''Chaetodon striatus''
**''Chaetodon semeion''
**''Chaetodon ephippium''
**''Chaetodon selene''
**''Chaetodon melannotus''
**''Chaetodon ocellicaudus''
**''Chaetodon rafflesi''
**''Chaetodon auriga''
**''Chaetodon vagabundus''
**''Chaetodon collare''
**''Chaetodon adiergastos''
**''Chaetodon fasciatus''
**''Chaetodon lunula''
**''Chaetodon semilarvatus''
**''Chaetodon lineolatus''
**''Chaetodon oxycephalus''
**''Chaetodon falcula''
**''Chaetodon ulietensis''

Evolution & fossil record

The earliest confirmed specimens of ''Chaetodon'' are from Early Oligocene-aged coral reef deposits of Italy, belonging to the extinct species ''Chaetodon wattsi''. This fossil is roughly concurrent when the family is thought to have radiated into its modern genus diversity. Indeterminate ''Chaetodon'' fossil specimens (including both articulated specimens and otoliths) referred to '' Chaetodon hoefleri'' are known from the Late Oligocene of Slovenia and the mid-late Miocene of Poland.

As it is not easy to distinguish this genus from close relatives, it may be that some Oligocene fossils are actually of other Chaetodontidae. The fossil record of this family is scant, with only ''Chelmon'' (or some similar genus like '' Chelmonops'' or '' Coradion'') being known from Miocene remains. As even crude molecular clocks suggest that the ancestors of the ''Chaetodon'' and the ''Chelmops'' lineage diverged in the Late Eocene already, nothing more can be said without new fossils being discovered.Paleobiology Database
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Footnotes

References

* Blum, S. D. (1988): ''Osteology and Phylogeny of the Chaetodontidae (Pisces: Perciformes)''. University of Hawaii, Honolulu.
* Fessler, Jennifer L. & Westneat, Mark W. (2007): Molecular phylogenetics of the butterflyfishes (Chaetodontidae): Taxonomy and biogeography of a global coral reef fish family. ''Molecular Phylogenetics and Evolution'' 45(1): 50–68.
* Hsu, Kui-Ching; Chen, Jeng-Ping & Shao, Kwang-Tsao (2007): Molecular phylogeny of ''Chaetodon'' (Teleostei: Chaetodontidae) in the Indo-West Pacific: evolution in geminate species pairs and species groups. ''Raffles Bulletin of Zoology'' Supplement 14: 77-86
PDF fulltext
* Sepkoski, Jack (2002): 'Chaetodon'' ''In:'' A compendium of fossil marine animal genera. ''Bulletins of American Paleontology'' 364: 560
HTML database excerpt

External links

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Taxa named by Carl Linnaeus
Marine fish genera
Rupelian genus first appearances
Extant Rupelian first appearances