Parrotfish (named for their mouths, which resemble a parrot's beak) are a clade of fish placed in the tribe Scarini of the

wrasse The wrasses are a family, Labridae, of marine ray-finned fish, many of which are brightly colored. The family is large and diverse, with over 600 species in 81 genera, which are divided into nine subgroups or tribes. They are typically small, ...

family (Labridae). Traditionally treated as their own family (Scaridae), genetic studies have found them to be deeply nested within the wrasses, and they are now treated as a subfamily (Scarinae) or tribe (Scarini) of them. With roughly 95 species, this group's largest species richness is in the

Indo-Pacific The Indo-Pacific is a vast biogeographic region of Earth. In a narrow sense, sometimes known as the Indo-West Pacific or Indo-Pacific Asia, it comprises the tropical waters of the Indian Ocean, the western and central Pacific Ocean, and the ...

. They are found in

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, rocky coasts, and

seagrass Seagrasses are the only flowering plants which grow in marine (ocean), marine environments. There are about 60 species of fully marine seagrasses which belong to four Family (biology), families (Posidoniaceae, Zosteraceae, Hydrocharitaceae and ...

beds, and can play a significant role in bioerosion.

Taxonomy

Traditionally, the parrotfishes have been considered to be a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

level

taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

, Scaridae. Although

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

and evolutionary analyses of parrotfishes are ongoing, they are now accepted to be a

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 ...

in the wrasses closely related to the

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 ...

Cheilini, and are now commonly referred to as scarine labrids (tribe Scarini, family Labridae). Some authorities have preferred to maintain the parrotfishes as a family-level taxon,Randall, J. E. (2007). ''Reef and Shore Fishes of the Hawaiian Islands.'' resulting in Labridae not being

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

(unless split into several families). The following taxonomic placement is based on Bellwood ''et al'' (2019): * Tribe Scarini ** Subtribe Scarina *** genus '' Bolbometopon'' Smith, 1956 (1 species) *** genus '' Cetoscarus'' Smith, 1956 (2 species) *** genus '' Chlorurus'' Swainson, 1839 (18 species) *** genus '' Hipposcarus'' Smith, 1956 (2 species) *** genus ''

Scarus ''Scarus'' is a genus of parrotfishes. With 52 currently recognised Extant taxon, extant species,Froese, R. and D. Pauly, Eds''Scarus''.FishBase. 2013. it is by far the largest parrotfish genus. The vast majority are found at coral reef, reefs i ...

'' Forsskål, 1775 (53 species) ** Subtribe Sparisomatina *** genus '' Calotomus'' Gilbert, 1890 (5 species) *** genus '' Cryptotomus'' Cope, 1870 (1 species) *** genus '' Leptoscarus'' Swainson, 1839 (1 species) *** genus '' Nicholsina'' Fowler, 1915 (3 species) *** genus '' Sparisoma'' Swainson, 1839 (15 species) Some sources retain the Scaridae as a family, placing it alongside the

s of the family Labridae and the weed whitings Odacidae in the order

Labriformes Labriformes is an Order (biology), order of ray-finned fishes within the clade Percomorpha. Some authors include the Labriformes as the clade Labroidei within the Perciformes while others include more Family (biology), families within the Labrif ...

, part of the Percomorpha. They also do not support the division of the Scaridae into two subfamilies. However, as such a placement is

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 ...

, they are placed within the wrasses by ''

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 ...

''. Fossil remains of parrotfishes are known dating back to the

Early Miocene The Early Miocene (also known as Lower Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages: the Aquitanian age, Aquitanian and Burdigalian stages. The sub-epoch lasted from 23.03 ± 0.05 annum, Ma to ...

Java Java is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea (a part of Pacific Ocean) to the north. With a population of 156.9 million people (including Madura) in mid 2024, proje ...

, Indonesia.

Description

Parrotfish are named for their

dentition Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiology ...

, which is distinct from other fish, including other labrids. Their numerous teeth are arranged in a tightly packed mosaic on the external surface of their jaw bones, forming a

parrot Parrots (Psittaciformes), also known as psittacines (), are birds with a strong curved beak, upright stance, and clawed feet. They are classified in four families that contain roughly 410 species in 101 genus (biology), genera, found mostly in ...

-like beak with which they rasp

algae Algae ( , ; : alga ) is an informal term for any organisms of a large and diverse group of photosynthesis, photosynthetic organisms that are not plants, and includes species from multiple distinct clades. Such organisms range from unicellular ...

from coral and other rocky substrates (which contributes to the process of bioerosion). Maximum sizes vary within the group, with the majority of species reaching in length. However, a few species reach lengths in excess of , and the green humphead parrotfish can reach up to . The smallest species is the bluelip parrotfish (''Cryptotomus roseus''), which has a maximum size of .

Protective mucus

Some parrotfish species, such as the queen parrotfish (''Scarus vetula''), secrete a mucus cocoon, particularly at night.Cerny-Chipman, E.
Distribution of Ultraviolet-Absorbing Sunscreen Compounds Across the Body Surface of Two Species of Scaridae.

DigitalCollections@SIT
' 2007. Accessed 2009-06-21. Prior to going to sleep, some species extrude mucus from their mouths, forming a protective cocoon that envelops the fish, presumably hiding its scent from potential predators. This mucus envelope may also act as an early warning system, allowing the parrotfish to flee when it detects predators such as

moray eel Moray eels, or Muraenidae (), are a family (biology), family of eels whose members are found worldwide. There are approximately 200 species in 15 genera which are almost exclusively Marine (ocean), marine, but several species are regu ...

s disturbing the membrane. The skin itself is covered in another mucous substance which may have antioxidant properties helpful in repairing bodily damage, or repelling parasites, in addition to providing protection from UV light.

Feeding

Most parrotfish species are

herbivore A herbivore is an animal anatomically and physiologically evolved to feed on plants, especially upon vascular tissues such as foliage, fruits or seeds, as the main component of its diet. These more broadly also encompass animals that eat ...

s, feeding mainly on epilithic algae.Bonaldo, R.M. & R.D. Rotjan (2018). The Good, the Bad, and the Ugly: Parrotfishes as Coral Predators. ''in'' Hoey, A.S. & R.M. Bonaldo, eds. Biology of Parrotfishes. CRC Press. A wide range of other small organisms are sometimes eaten, including invertebrates ( sessile and

benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...

species, as well as

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 ...

), bacteria and

detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...

. A few mostly larger species such as the green humphead parrotfish (''Bolbometopon muricatum'') feed extensively on living

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). None of these are exclusive corallivores, but polyps can make up as much as half their diet or even more in the green humphead parrotfish. Overall it has been estimated that fewer than one percent of parrotfish bites involve live corals and all except the green humphead parrotfish prefer algae-covered surfaces over live corals. Nevertheless, when they do eat coral polyps, localized coral death can occur. Their feeding activity is important for the production and distribution of coral sands in the reef

biome A biome () is a distinct geographical region with specific climate, vegetation, and animal life. It consists of a biological community that has formed in response to its physical environment and regional climate. In 1935, Tansley added the ...

, and can prevent algal overgrowth of the reef structure. The teeth grow continuously, replacing material worn away by feeding. Whether they feed on coral, rock or seagrasses, the substrate is ground up between the pharyngeal teeth. After they digest the edible portions from the rock, they excrete it as sand, helping create small islands and the sandy beaches. The humphead parrotfish can produce of sand each year. Or, on average (as there are so many variables i.e. size/species/location/depth etc.), almost per parrotfish per day. While feeding, parrotfish must be cognizant of predation by one of their main predators, the

lemon shark The lemon shark (''Negaprion brevirostris'') is a species of shark from the family Requiem shark, Carcharhinidae, known for its yellowish skin, which inspires its common name. It is classified as a Vulnerable species by the International Union for ...

. On Caribbean coral reefs, parrotfish are important consumers of

sponge Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a basal clade and a sister taxon of the diploblasts. They are sessile filter feeders that are bound to the seabed, and a ...

s. An indirect effect of parrotfish grazing on sponges is the protection of reef-building corals that would otherwise be overgrown by fast-growing sponge species. Analysis of parrotfish feeding biology describes three functional groups: excavators, scrapers and browsers. Excavators have larger, stronger jaws that can gouge the substrate, leaving visible scars on the surface. Scrapers have less powerful jaws that can but infrequently do leave visible scraping scars on the substrate. Some of these may also feed on sand instead of hard surfaces. Browsers mainly feed on seagrasses and their

epiphyte An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...

s. Mature excavating species include '' Bolbometopon muricatum'', '' Cetoscarus'', '' Chlorurus'' and '' Sparisoma viride''. These excavating species all feed as scrapers in early juvenile stages, but '' Hipposcarus'' and ''

'', which also feed as scrapers in early juvenile stages, retain the scraping feeding mode as adults. Browsing species are found in the genera '' Calotomus'', '' Cryptotomus'', '' Leptoscarus'', '' Nicholsina'' and ''Sparisoma''. Feeding modes reflect habitat preferences, with browsers chiefly living in the grassy seabed, and excavators and scrapers on coral reefs. Recently, the microphage feeding hypothesis challenged the prevailing paradigm of parrotfish as algal consumers by proposing that: Microscopy and molecular barcoding of coral reef substrate bitten by scraping and excavating parrotfish suggest that coral reef cyanobacteria from the order Nostocales are important in the feeding of these parrotfish. Additional microscopy and molecular barcoding research indicates that some parrotfish may ingest microscopic biota associated with endolithic sponges.

Life cycle

Most tropical species form large schools when feeding and these are often grouped by size. Harems of several females presided over by a single male are normal in most species, with the males vigorously defending their position from any challenge. As pelagic spawners, parrotfish release many tiny, 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 ...

. The eggs float freely, settling into the coral until hatching.

Sex change

The development of parrotfishes is complex and accompanied by a series of changes in sex and colour ( polychromatism). Most species are sequential hermaphrodites, starting as females (known as the initial phase) and then changing to males (the terminal phase). In many species, for example the stoplight parrotfish (''Sparisoma viride''), a number of individuals develop directly to males (i.e., they do not start as females). These directly developing males usually most resemble the initial phase, and often display a different mating strategy than the terminal phase males of the same species.Bester, C.
Stoplight parrotfish.
'' Florida Museum of Natural History, Ichthyology Department. Accessed 15-12-2009 A few species such as the Mediterranean parrotfish (''S. cretense'') are secondary gonochorists. This means that some females do not change sex (they remain females throughout their lives), the ones that do change from female to male do it while still immature (reproductively functioning females do not change to males) and there are no males with female-like colors (the initial phase males in other parrotfish). The marbled parrotfish (''Leptoscarus vaigiensis'') is the only species of parrotfish known not to change sex. In most species, the initial phase is dull red, brown, or grey, while the terminal phase is vividly green or blue with bright pink, orange or yellow patches. In a smaller number of species the phases are similar, and in the Mediterranean parrotfish the adult female is brightly colored, while the adult male is gray. In most species, juveniles have a different color pattern from adults. Juveniles of some tropical species can alter their color temporarily to

mimic In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...

other species. Where the sexes and ages differ, the remarkably different phases often were first described as separate species. As a consequence early scientists recognized more than 350 parrotfish species, which is almost four times the actual number. The sex change in parrotfishes is accompanied by changes in circulating steroids. Females have high levels of estradiol, moderate levels of T and undetectable levels of the major fish androgen 11-ketotestosterone. During the transition from initial to terminal coloration phases, concentrations of 11-ketotestosterone rise dramatically and estrogen levels decline. If a female is injected with 11-ketotestosterone, it will cause a precocious change in gonadal, gametic and behavioural sex.

Economic importance

A commercial

fishery Fishery can mean either the enterprise of raising or harvesting fish and other aquatic life or, more commonly, the site where such enterprise takes place ( a.k.a., fishing grounds). Commercial fisheries include wild fisheries and fish far ...

exists for some of the larger species, particularly in the Indo-Pacific, but also for a few others like the Mediterranean parrotfish. Protecting parrotfishes is proposed as a way of saving Caribbean coral reefs from being overgrown with seaweed and sponges. Despite their striking colors, their feeding behavior renders them highly unsuitable for most marine aquaria. A new study has discovered that the parrotfish is extremely important for the health of the

Great Barrier Reef The Great Barrier Reef is the world's largest coral reef system, composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...

; it is the only one of thousands of reef fish species that regularly performs the task of scraping and cleaning inshore coral reefs.

Gallery

File:Scarus globiceps mâle.jpg, '' Scarus globiceps'' (male) File:Parrotfish turquoisse.jpg, '' Chlorurus microrhinos'' File:Bolbometopon muricatum.jpg, '' Bolbometopon muricatum'' File:Viridescent Parrotfish - Calotomus viridescens.jpg, '' Calotomus viridescens'' File:Cetoscarus ocellatus Great Barrier Reef.jpg, '' Cetoscarus ocellatus'' File:Chlorurus sordidus by Jaroslaw Barski.jpg, '' Chlorurus sordidus'' File:Hipposcarus longiceps.jpg, '' Hipposcarus longiceps'' File:Queen parrotfish Scarus vetula (2442375123).jpg, '' Scarus vetula'' File:Stoplight-parrotfish.jpg, '' Sparisoma viride''

Timeline of genera

ImageSize = width:1000px height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-65.5 till:10 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:5 start:-65.5 ScaleMinor = unit:year increment:1 start:-65.5 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: -65.5 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:

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 ...

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: -65.5 till: -23.03 color:paleogene text:

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 ...

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: -33.9 till: 0 text:

PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 bar:period from: -65.5 till: -55.8 color:paleocene text: