The butterflyfish are a group of conspicuous

tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

marine

fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...

of the family Chaetodontidae; the bannerfish and coralfish are also included in this group. The approximately 129

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

in 12 genera are found mostly on the

reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock o ...

s of the

Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the " Old World" of Africa, Europe an ...

, Indian, and

Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contin ...

s. A number of

species pair In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

s occur in the Indian and Pacific Oceans, members of the huge genus '' Chaetodon''. Butterflyfish look like smaller versions of angelfish (

Pomacanthidae Marine angelfish are perciform fish of the family Pomacanthidae. They are found on shallow reefs in the tropical Atlantic, Indian, and mostly western Pacific Oceans. The family contains seven genera and about 86 species. They should not be conf ...

), but unlike these, lack

preopercle The operculum is a series of bones found in bony fish and chimaeras that serves as a facial support structure and a protective covering for the gills; it is also used for respiration and feeding. Anatomy The opercular series contains four bon ...

spines at the

gill cover The operculum is a series of bones found in bony fish and chimaeras that serves as a facial support structure and a protective covering for the gills; it is also used for respiration and feeding. Anatomy The opercular series contains four bon ...

s. Some members of the genus ''

Heniochus ''Heniochus'' is a genus of marine ray-finned fish, butterflyfishes from the family Chaetodontidae. They are native to the Indo-Pacific. Though very similar in appearance to the Moorish idol ''( Zanclus cornutus)'', the members of this genus ar ...

'' resemble the

Moorish idol The Moorish idol (''Zanclus cornutus'') is a marine fish species, the sole extant representative of the family Zanclidae (from the Greek ζαγκίος, ''zagkios'', "oblique") in order Acanthuriformes. A common inhabitant of tropical to subtr ...

(''Zanclus cornutus'') of the

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...

Zanclidae. Among the paraphyletic

Perciformes Perciformes (), also called the Percomorpha or Acanthopteri, is an order or superorder of ray-finned fish. If considered a single order, they are the most numerous order of vertebrates, containing about 41% of all bony fish. Perciformes means ...

, the former are probably not too distantly related to butterflyfish, whereas the Zanclidae seem far less close.

Description and ecology

Butterflyfish mostly range from in length. The largest species, the

lined butterflyfish The lined butterflyfish (''Chaetodon lineolatus'') is a species of marine ray-finned fish. a butterflyfish belonging to the family Chaetodontidae, one of the largest species in the genus ''Chaetodon''. It has a wide range from the Red Sea to Sou ...

and the

saddle butterflyfish The saddle butterflyfish (''Chaetodon ephippium'') is a species of marine ray-finned fish, a butterflyfish belonging to the family Chaetodontidae. It is found in the Indian and Pacific Oceans from Sri Lanka and the Cocos-Keeling Islands to the ...

, ''C. ephippium'', grow to . The common name references the brightly coloured and strikingly patterned bodies of many species, bearing shades of black, white, blue, red, orange, and yellow. Other species are dull in colour. Butterflyfish are a boundless, different group of marine perciods with delegates on practically all coral reef frameworks and in every single tropical ocean. Their bright and color patterns have drawn in much consideration, creating an abundance of data about their conduct and environment. Many have eyespots on their flanks and dark bands across their eyes, not unlike the patterns seen on

butterfly Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprise ...

wings. Their deep,

lateral Lateral is a geometric term of location which may refer to: Healthcare *Lateral (anatomy), an anatomical direction * Lateral cricoarytenoid muscle * Lateral release (surgery), a surgical procedure on the side of a kneecap Phonetics *Lateral co ...

ly narrow bodies are easily noticed through the profusion of reef life. The conspicuous coloration of butterflyfish may be intended for interspecies communication. Butterflyfish have uninterrupted

dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through c ...

s with tail fins that may be rounded or truncated, but are never forked. Generally diurnal and frequenting waters less than deep (though some species descend to , butterflyfish stick to particular home ranges. These

corallivore A corallivore is an animal that feeds on coral. Corallivores are an important group of reef organism because they can influence coral abundance, distribution, and community structure. Corallivores feed on coral using a variety of unique adaptati ...

s are especially territorial, forming pairs and staking claim to a specific coral head. Contrastingly, the

zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...

feeders form large conspecific groups. By night, butterflyfish hide in reef crevices and exhibit markedly different coloration. Their coloration also makes them popular aquarium fish. However, most species feed on

coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and ...

polyps and sea anemones. Balancing the relative populations of prey and predator is complex, leading hobby aquarists to focus on the few generalists and specialist

feeders. Butterflyfish are pelagic spawners; that is, they release many buoyant eggs into the water, which become part of the

plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...

, floating with the currents until hatching. The fry go through a tholichthys stage, wherein the body of the post

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...

l fish is covered in large, bony plates extending from the head. They lose their bony plates as they mature. Only one other family of fish, the scats ( Scatophagidae) express such an armored stage.

Taxonomy, systematics and evolution

The Chaetodontidae can be, but are not usually, divided into two lineages that arguably are subfamilies. The subfamily name Chaetodontinae is a little-used leftover from the period when the

and Chaetodontidae were united under the latter name as a single family. Hence, Chaetodontinae is today considered a junior synonym of Chaetodontidae. In any case, one lineage of Chaetodontidae (in the modern sense) contains the "typical" butterflyfish around ''Chaetodon'', while the other unites the bannerfish and coralfish genera. As the

are highly paraphyletic, the precise relationships of the Chaetodontidae as a whole are badly resolved.Fessler & Westneat (2007) Chaetodontidae is classified within the suborder

Percoidei Percoidei is one of 3 suborders of bony fishes in the order Perciformes. Many commercially harvested fish species are considered to be contained in this suborder, including the snappers, groupers, basses, goatfishes and perches. Divisions The ...

by the 5th edition of '' Fishes of the World'', but they are placed in an unnamed clade which sits outside the superfamily

Percoidea Percoidea is a superfamily of fish of the order Perciformes. The superfamily includes about 3,374 species. Classification The Percoidesa are classified in the 5th Edition of the ''Fishes of the World'' as follows: * Percoidea * ...

. This clade contains 7 families which appear to have some relationship to Acanthuroidei, Monodactylidae, and

Priacanthidae The Priacanthidae, the bigeyes, are a family of 18 species of marine ray-finned fishes. " Catalufa" is an alternate common name for some members of the Priacanthidae. The etymology of the scientific name (, to bite + , thorn) refers to the fam ...

. Other authorities have paced the family in the order Chaetodontiformes alongside the family

Leiognathidae Leiognathidae, the ponyfishes, slipmouths or slimys / slimies, are a small family of fishes in the order Perciformes. They inhabit marine and brackish waters in the Indian and West Pacific Oceans. They can be used in the preparation of ''bagoong' ...

. Before DNA sequencing, the

taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...

was confused about whether to treat these as species or subspecies. Also, numerous subgenera have been proposed for splitting out of ''Chaetodon'', and it is becoming clear how to subdivide the genus if that is desired. The

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

record of this group is marginal. Their restriction to

coral reef A coral reef is an underwater ecosystem characterized by reef-building corals. Reefs are formed of colonies of coral polyps held together by calcium carbonate. Most coral reefs are built from stony corals, whose polyps cluster in groups. C ...

s means their carcasses are liable to be dispersed by scavengers, overgrown by

s, and any that do fossilize will not long survive

erosion Erosion is the action of surface processes (such as water flow or wind) that removes soil, rock, or dissolved material from one location on the Earth's crust, and then transports it to another location where it is deposited. Erosion is dis ...

. However, '' Pygaeus'', a very basal fossil from the mid- to late

Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...

, dates from around the

Bartonian The Bartonian is, in the ICS's geologic time scale, a stage or age in the middle Eocene Epoch or Series. The Bartonian Age spans the time between . It is preceded by the Lutetian and is followed by the Priabonian Age. Stratigraphic defini ...

40–37 million years ago (Mya). Thus, the Chaetodontidae emerged probably in the early to mid-Eocene. A crude

molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...

in combination with the evidence given by ''Pygaeus'' allows placement of the initial split between the two main lineages to the middle to late Eocene, and together with the few other fossils, it allows the deduction that most living genera were probably distinct by the end of the Paleogene 23 Mya.

Genera

The bannerfish-coralfish lineage can be further divided in two groups; these might be considered

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confli ...

s, but have not been formally named. Genera are listed in order of the presumed

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...

, from the most ancient to the youngest: Bannerfish/coralfish lineage 1: * '' Amphichaetodon'' Burgess, 1978 * '' Coradion'' Kaup, 1860 * '' Chelmon'' Cloquet, 1817 * '' Chelmonops'' Bleeker, 1876 Bannerfish/coralfish lineage 2: * '' Forcipiger''

Jordan Jordan ( ar, الأردن; tr. ' ), officially the Hashemite Kingdom of Jordan,; tr. ' is a country in Western Asia. It is situated at the crossroads of Asia, Africa, and Europe, within the Levant region, on the East Bank of the Jordan Rive ...

McGregor McGregor may refer to: People * McGregor (surname) * Clan MacGregor, a Scottish highland clan * McGregor W. Scott (born 1962), U.S. attorney Characters * Mr. McGregor, a fictional character from Peter Rabbit Places in Canada: * McGregor Lake ...

, 1898 * '' Hemitaurichthys'' Bleeker, 1876 * ''

'' Cuvier, 1816 * '' Johnrandallia'' Nalbant, 1974 The "typical" butterflyfishes may eventually come to contain more genera; see '' Chaetodon'': * '' Chaetodon''

Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...

, 1758 * '' Parachaetodon'' Bleeker, 1874 * ''

Prognathodes ''Prognathodes'' is a genus of butterflyfish in the family Chaetodontidae. They are found at rocky reefs in tropical oceans around the world and are mainly found deeper than ''Chaetodon''; often a depths below those possible in normal scuba divi ...

Gill A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...

, 1862 * '' Roa'' Jordan, 1923

Timeline

ImageSize = width:1000px height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-66 till:10 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:5 start:-66 ScaleMinor = unit:year increment:1 start:-66 TimeAxis = orientation:hor AlignBars = justify Colors = #legends id:CAR value:claret id:ANK value:rgb(0.4,0.3,0.196) id:HER value:teal id:HAD value:green id:OMN value:blue id:black value:black id:white value:white id:cenozoic value:rgb(0.54,0.54,0.258) id:paleogene value:rgb(0.99,0.6,0.32) id:paleocene value:rgb(0.99,0.65,0.37) id:eocene value:rgb(0.99,0.71,0.42) id:oligocene value:rgb(0.99,0.75,0.48) id:neogene value:rgb(0.999999,0.9,0.1) id:miocene value:rgb(0.999999,0.999999,0) id:pliocene value:rgb(0.97,0.98,0.68) id:quaternary value:rgb(0.98,0.98,0.5) id:pleistocene value:rgb(0.999999,0.95,0.68) id:holocene value:rgb(0.999,0.95,0.88) BarData= bar:eratop bar:space bar:periodtop bar:space bar:NAM1 bar:NAM2 bar:space bar:period bar:space bar:era PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(7,-4) bar:periodtop from: -66 till: -55.8 color:paleocene text:

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...

from: -55.8 till: -33.9 color:eocene text:

from: -33.9 till: -23.03 color:oligocene text: Oligocene from: -23.03 till: -5.332 color:miocene text:

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

from: -5.332 till: -2.588 color:pliocene text: Plio. from: -2.588 till: -0.0117 color:pleistocene text: Pleist. from: -0.0117 till: 0 color:holocene text: H. bar:eratop from: -66 till: -23.03 color:paleogene text: Paleogene from: -23.03 till: -2.588 color:neogene text: Neogene from: -2.588 till: 0 color:quaternary text: Q. PlotData= align:left fontsize:M mark:(line,white) width:5 anchor:till align:left color:oligocene bar:NAM1 from: -28.4 till: 0 text: Chaetodon color:miocene bar:NAM2 from: -23.03 till: 0 text: Chelmon PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 bar:period from: -66 till: -55.8 color:paleocene text:

from: -55.8 till: -33.9 color:eocene text:

from: -33.9 till: -23.03 color:oligocene text: Oligocene from: -23.03 till: -5.332 color:miocene text:

from: -5.332 till: -2.588 color:pliocene text: Plio. from: -2.588 till: -0.0117 color:pleistocene text: Pleist. from: -0.0117 till: 0 color:holocene text: H. bar:era from: -66 till: -23.03 color:paleogene text: Paleogene from: -23.03 till: -2.588 color:neogene text: Neogene from: -2.588 till: 0 color:quaternary text: Q.

Gallery

Image:Seattle Aquarium, 1.JPG, Copperband butterflyfish, ''Chelmon rostratus'' Image:Johnrandallia nigrirostris.jpg, The enigmatic '' Johnrandallia nigrirostris'' Image:Chaetodon kleinii.jpg, Sunburst butterflyfish, (sometimes placed in ''Lepidochaetodon'') Image:Bep chaetodon bennetti.jpg, Bluelashed butterflyfish, ''Chaetodon bennetti (sometimes placed in ''Megaprotodon'')

References

;General * (2007): Molecular phylogenetics of the butterflyfishes (Chaetodontidae): Taxonomy and biogeography of a global coral reef fish family. '' Mol. Phylogenet. Evol.'' 45(1): 50–68. (HTML abstract) * 008br>Family Chaetodontidae – Butterflyfishes
Retrieved 2008-SEP-02. * (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
* (2002): 'Chaetodon'' ''In:'' A compendium of fossil marine animal genera. ''Bulletins of American Paleontology'' 364: 560
HTML database excerpt
;Specific {{Taxonbar, from=Q271004 Articles which contain graphical timelines Extant Maastrichtian first appearances Taxa named by Constantine Samuel Rafinesque