The butterflyfish are a group of conspicuous

tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

marine

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

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

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

in 12

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

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 component, abiotic (non-living) processes such as deposition (geol ...

s of the

Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...

, Indian, and

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

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 ''Chaetodon'' is a tropical fish genus in the family (biology), 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, ...

''. 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 co ...

), 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 Chimaera, 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 contain ...

s. Some members of the genus '' Heniochus'' resemble the Moorish idol (''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 "unisp ...

Zanclidae. Among the

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

Perciformes Perciformes (), also called the Acanthopteri, is an order or superorder of ray-finned fish in the clade Percomorpha. ''Perciformes'' means " perch-like". Among the well-known members of this group are perches and darters ( Percidae), and als ...

, 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 th ...

, ''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 percoids 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 winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

wings. Their deep,

lateral Lateral is a geometric term of location which may also refer to: Biology and healthcare * Lateral (anatomy), a term of location meaning "towards the side" * Lateral cricoarytenoid muscle, an intrinsic muscle of the larynx * Lateral release ( ...

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 on the back of most marine and freshwater vertebrates. Dorsal fins have evolved independently several times through convergent evolution adapting to marine environments, so the fins are not all homologous. They are found ...

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

zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...

feeders form large conspecific groups. By night, butterflyfish hide in reef crevices and exhibit markedly different coloration. Their coloration also makes them popular

aquarium An aquarium (: aquariums or aquaria) is a vivarium of any size having at least one transparent side in which aquatic plants or animals are kept and displayed. fishkeeping, Fishkeepers use aquaria to keep fish, invertebrates, amphibians, aquati ...

fish. However, most species feed on

coral Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact Colony (biology), colonies of many identical individual polyp (zoology), polyps. Coral species include the important Coral ...

polyps and

sea anemone Sea anemones ( ) are a group of predation, predatory marine invertebrates constituting the order (biology), order Actiniaria. Because of their colourful appearance, they are named after the ''Anemone'', a terrestrial flowering plant. Sea anemone ...

s. 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 that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...

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

larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...

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 In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...

. 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 In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

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

. As the

are highly

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

suborder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and recognized ...

Percoidei Percoidei is a suborder of bony fishes in the order Perciformes. Many commercially harvested fish species are considered to be contained in this suborder, including the groupers, Serranidae, seabasses and perches. Divisions The following classif ...

by the 5th edition of ''

Fishes of the World ''Fishes of the World'' is a standard reference for the systematics of fishes. It was first written in 1976 by the American ichthyologist Joseph S. Nelson (1937–2011). Now in its fifth edition (2016), the work is a comprehensive overview of t ...

'', but they are placed in an unnamed

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

which sits outside the superfamily

Percoidea Percomorpha () is an extremely large and diverse clade of ray-finned fish. With more than 17,000 known species (including tuna, seahorses, gobies, cichlids, flatfish, wrasse, perches, anglerfish, and pufferfish) known from both marine and fresh ...

. This clade contains 7 families which appear to have some relationship to

Acanthuroidei Acanthuroidei , is a group of ray finned fishes which is a suborder of the Acanthuriformes, although older classifications regarded it as a suborder of the Perciformes, the largest order (biology), order of fish, The suborder includes the surgeonf ...

, Monodactylidae, and

Priacanthidae The Priacanthidae, the bigeyes, are a Family (biology), family of 18 species of marine ray-finned fishes. "Catalufa (disambiguation), Catalufa" is an alternate common name for some members of the Priacanthidae. The etymology of the scientific ...

. Other authorities have paced the family in the order Chaetodontiformes alongside the family Leiognathidae. Presently, based on the most recent phylogenetic analyses,

Eschmeyer's Catalog of Fishes Catalog of Fishes is a comprehensive on-line database and reference work on the scientific names of fish species and genera. It is global in its scope and is hosted by the California Academy of Sciences. It has been compiled and is continuously up ...

places it in the large, diverse order

Acanthuriformes Acanthuriformes is a large, diverse order of mostly marine ray-finned fishes, part of the Percomorpha clade. In the past, members of this clade were placed in the suborders Acanthuroidea and Percoidea of the order Perciformes, but this treatment ...

. Molecular analyses suggest that the Chaetodontidae shared a common ancestor with the Scatophagidae and

that lived during the

Early Eocene In the geologic timescale the Ypresian is the oldest age (geology), age or lowest stage (stratigraphy), stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by th ...

(54 Ma). Before

DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...

, the

taxonomy image:Hierarchical clustering diagram.png, 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme o ...

was confused about whether to treat these as species or

subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...

. Also, numerous

subgenera In biology, a subgenus ( subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between the ge ...

have been proposed for splitting out of ''Chaetodon'', and it is becoming clear how to subdivide the genus if that is desired.

Fossil record

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 preserve ...

record of this group is disputed and 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 group ...

s means their carcasses are liable to be dispersed by

scavenger Scavengers are animals that consume Corpse decomposition, dead organisms that have died from causes other than predation or have been killed by other predators. While scavenging generally refers to carnivores feeding on carrion, it is also a he ...

s, overgrown by

s, and any that do fossilize will not long survive

erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...

. Many

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

-aged fish such as '' Pygaeus'' have been variously referred to Chaetodontidae in the past; if used for calibration, it can be deduced that most living genera were probably distinct by the end of the

Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...

23 Mya. However, these Eocene genera cannot be reliably referred to Chaetodontidae. The earliest confirmed fossil specimens of Chaetodontidae are an indeterminate larval specimen from Frauenweiler, Germany, two tholichthys specimens from the Menilite Formation of Poland, as well as the extinct species '' Chaetodon (Blumchaetodon) wattsi'' from Italy; all date to the

Early Oligocene The Rupelian is, in the geologic timescale, the older of two age (geology), ages or the lower of two stage (stratigraphy), stages of the Oligocene epoch (geology), Epoch/series (stratigraphy), Series. It spans the time between . It is preceded b ...

. The occurrence of the first confirmed butterflyfish in the Early Oligocene is supported by molecular phylogenies indicating that the family may have arose during the

Late Eocene The Priabonian is, in the International Commission on Stratigraphy, ICS's geologic timescale, the latest age (geology), age or the upper stage (stratigraphy), stage of the Eocene epoch (geology), Epoch or series (stratigraphy), Series. It spans ...

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 use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

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

phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...

, from the most ancient to the youngest: Bannerfish/coralfish lineage 1: * '' Amphichaetodon'' Burgess, 1978 * '' Coradion'' Kaup, 1860 * ''

Chelmon ''Chelmon'' is a genus of marine ray-finned fish in the family Chaetodontidae, the butterflyfishes. They are tropical species native to the western Pacific Ocean. Species There are currently three recognized species in this genus: * '' Chelmon ...

'' Cloquet, 1817 * '' Chelmonops'' Bleeker, 1876 Bannerfish/coralfish lineage 2: * ''

Forcipiger ''Forcipiger'' is a genus of fish in the family butterflyfish, Chaetodontidae, the butterflyfishes. It is distributed throughout the Indo-Pacific region. The name of this genus means “bearing forceps” and is a reference to the long, slender s ...

Jordan Jordan, officially the Hashemite Kingdom of Jordan, is a country in the Southern Levant region of West Asia. Jordan is bordered by Syria to the north, Iraq to the east, Saudi Arabia to the south, and Israel and the occupied Palestinian ter ...

& McGregor, 1898 * '' Hemitaurichthys'' Bleeker, 1876 * '' Heniochus''

Cuvier Jean Léopold Nicolas Frédéric, baron Cuvier (23 August 1769 – 13 May 1832), known as Georges Cuvier (; ), was a French natural history, naturalist and zoology, zoologist, sometimes referred to as the "founding father of paleontology". Cuv ...

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

'': * ''

Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

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

Gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...

, 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 (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...

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

from: -33.9 till: -23.03 color:oligocene text:

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

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:

from: -23.03 till: -2.588 color:neogene text:

Neogene The Neogene ( ,) is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period million years ago. It is the second period of th ...

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:

color:miocene bar:NAM2 from: -23.03 till: 0 text:

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:

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:

from: -23.03 till: -2.588 color:neogene text:

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) *

008 008, OO8, O08, or 0O8 may refer to: * "008", a fictional 00 Agent In Ian Fleming's James Bond novels and the derived films, the 00 Section of MI6 is considered the secret service's elite. A 00 (pronounced "Double O") is a field agent who ho ...

br>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 {{Authority control Articles which contain graphical timelines Taxa named by Constantine Samuel Rafinesque Extant Rupelian first appearances