Parrotfish#Economic importance

{{Short description|Clade of ray-finned fishes}}

{{Use dmy dates|date=April 2017}}

{{Redirect2|Scarinae|Scarini|the group of leafhoppers formerly known by these names|Gyponini}}

{{Distinguish|Parrot cichlid (disambiguation){{!}}parrot cichlid}}

{{Automatic taxobox

| fossil_range = {{fossil range|Miocene|present}}

| image = Scarus frenatus by Ewa Barska.jpg

| image_caption = Scarus frenatus

| taxon = Scarini

| authority = Rafinesque, 1810

| subdivision_ranks = Genera

| subdivision = * Bolbometopon

}}

Parrotfish (named for their mouths, which resemble a parrot's beak) are a clade of fish placed in the tribe Scarini of the wrasse family (Labridae).{{Cite journal |last1=Hughes |first1=Lily C |last2=Nash |first2=Chloe M |last3=White |first3=William T |last4=Westneat |first4=Mark W |date=2023-06-17 |editor-last=Matschiner |editor-first=Michael |title=Concordance and Discordance in the Phylogenomics of the Wrasses and Parrotfishes (Teleostei: Labridae) |url=https://academic.oup.com/sysbio/article/72/3/530/6798871 |journal=Systematic Biology |language=en |volume=72 |issue=3 |pages=530–543 |doi=10.1093/sysbio/syac072 |issn=1063-5157}} 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. They are found in coral reefs, rocky coasts, and seagrass beds, and can play a significant role in bioerosion.

Taxonomy

Traditionally, the parrotfishes have been considered to be a family level taxon, Scaridae. Although phylogenetic and evolutionary analyses of parrotfishes are ongoing, they are now accepted to be a clade in the wrasses closely related to the tribe Cheilini, and are now commonly referred to as scarine labrids (tribe Scarini, family Labridae).{{Cite journal |last1=Bellwood |first1=David R. |last2=Schultz |first2=Ortwin |last3=Siqueira |first3=Alexandre C. |last4=Cowman |first4=Peter F. |date=2019 |title=A review of the fossil record of the Labridae |url=https://www.jstor.org/stable/26595690 |journal=Annalen des Naturhistorischen Museums in Wien. Serie A für Mineralogie und Petrographie, Geologie und Paläontologie, Anthropologie und Prähistorie |volume=121 |pages=125–194 |jstor=26595690 |issn=0255-0091}} 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. {{ISBN|978-1-929054-03-9}} resulting in Labridae not being monophyletic (unless split into several families).

The following taxonomic placement is based on Bellwood et al (2019):

Some sources retain the Scaridae as a family, placing it alongside the wrasses of the family Labridae and the weed whitings Odacidae in the order Labriformes, part of the Percomorpha. They also do not support the division of the Scaridae into two subfamilies.{{cite book |author1=J. S. Nelson |url=https://sites.google.com/site/fotw5th/ |title=Fishes of the World |author2=T. C. Grande |author3=M. V. H. Wilson |publisher=Wiley |year=2016 |isbn=978-1-118-34233-6 |edition=5th |pages=429–430}} However, as such a placement is paraphyletic, they are placed within the wrasses by Eschmeyer's Catalog of Fishes.{{Cite web |last1=Fricke |first1=R. |last2=Eschmeyer |first2=W. N. |last3=Van der Laan |first3=R. |date=2025 |title=ESCHMEYER'S CATALOG OF FISHES: CLASSIFICATION |url=https://www.calacademy.org/eschmeyers-catalog-of-fishes-classification |access-date=2025-02-10 |website=California Academy of Sciences |language=en}}

Fossil remains of parrotfishes are known dating back to the Early Miocene of Java, Indonesia.

Description

File:Pseudoscarus.jpg

Parrotfish are named for their dentition,Ostéologie céphalique de deux poissons perroquets (Scaridae: Teleostei) TH Monod, JC Hureau, AE Bullock - Cybium, 1994 - Société française d'ichtyologie 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-like beak with which they rasp algae 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 {{convert|30|-|50|cm|abbr=on}} in length. However, a few species reach lengths in excess of {{convert|1|m|abbr=on}}, and the green humphead parrotfish can reach up to {{convert|1.3|m|abbr=on}}.{{FishBase |genus=Bolbometopon |species=muricatum |month=December |year=2009}} The smallest species is the bluelip parrotfish (Cryptotomus roseus), which has a maximum size of {{convert|13|cm|abbr=on}}.{{FishBase |genus=Cryptotomus |species=roseus |month=September |year=2015}}Shah, A.K. (2016). {{url|https://sta.uwi.edu/fst/lifesciences/sites/default/files/lifesciences/documents/ogatt/Cryptotomus_roseus - Slender Parrotfish.pdf|Cryptotomus roseus (Slender Parrotfish)}}. The Online Guide to the Animals of Trinidad and Tobago. The University of the West Indies. Accessed 11 March 2018.

Protective mucus

File:Scarus zelindae.jpg in its mucus cocoon|left]]

Some parrotfish species, such as the queen parrotfish (Scarus vetula), secrete a mucus cocoon, particularly at night.Cerny-Chipman, E. "[http://digitalcollections.sit.edu/cgi/viewcontent.cgi?article=1133&context=isp_collection Distribution of Ultraviolet-Absorbing Sunscreen Compounds Across the Body Surface of Two Species of Scaridae.]" [http://digitalcollections.sit.edu/ 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.Langerhans, R.B. "[http://faculty-staff.ou.edu/L/Randall.B.Langerhans-1/Langerhans%202006.pdf Evolutionary consequences of predation: avoidance, escape, reproduction, and diversification.] {{webarchive|url=https://web.archive.org/web/20110614031545/http://faculty-staff.ou.edu/L/Randall.B.Langerhans-1/Langerhans%202006.pdf |date=14 June 2011 }}" pp. 177–220 in Elewa, A.M.T. ed. Predation in organisms: a distinct phenomenon. Heidelberg, Germany, Springer-Verlag. 2007. Accessed 2009-06-21. This mucus envelope may also act as an early warning system, allowing the parrotfish to flee when it detects predators such as moray eels 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.

{{Clear|left}}

Feeding

File:Scaridae - Bolbometopon muricatum.jpg is suited to 'excavating', grinding the sturdiest corals.]]

File:BuDIha.jpg is suited to 'browsing' on seagrass, macroalgae, and epilithic algae without touching the rocky substrate.]]

Most parrotfish species are herbivores, feeding mainly on epilithic algae.{{Cite journal|last=Bellwood|first=David R.|date=1994-07-14|title=A phylogenetic study of the parrotfish family Scaridae (Pisces: Labroidea), with a revision of genera|journal=Records of the Australian Museum, Supplement|language=en|volume=20|pages=1–86|doi=10.3853/j.0812-7387.20.1994.51|issn=0812-7387|doi-access=free}}{{cite journal | last1 = Bellwood | first1 = D.R. | last2 = Choat | first2 = J.H. | year = 1990 | title = A functional analysis of grazing in parrotfishes (family Scaridae): the ecological implications | journal = Environ Biol Fish | volume = 28 | issue = 1–4| pages = 189–214 | doi = 10.1007/BF00751035 | s2cid = 11262999 }}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. {{isbn|978-1482224016}} A wide range of other small organisms are sometimes eaten, including invertebrates (sessile and benthic species, as well as zooplankton), bacteria and detritus.{{cite journal | last1 = Comeros-Raynal | first1 = Choat | last2 = Polidoro | first2 = Clements | last3 = Abesamis | first3 = Craig | last4 = Lazuardi | first4 = McIlwain | last5 = Muljadi | first5 = Myers | last6 = Nañola Jr | first6 = Pardede | last7 = Rocha | first7 = Russell | last8 = Sanciangco | first8 = Stockwell | last9 = Harwell | last10 = Carpenter | year = 2012 | title = The Likelihood of Extinction of Iconic and Dominant Herbivores and Detritivores of Coral Reefs: The Parrotfishes and Surgeonfishes | journal = PLOS ONE | volume = 7 | issue = 7| page = e39825 | doi = 10.1371/journal.pone.0039825 | pmid = 22808066 | pmc = 3394754 | bibcode = 2012PLoSO...739825C | doi-access = free }} A few mostly larger species such as the green humphead parrotfish (Bolbometopon muricatum) feed extensively on living 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, 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.{{cite book |title=Coral Reefs: Cities Under The Seas |last=Murphy |first=Richard C. |year=2002 |isbn=978-0-87850-138-0 |publisher=The Darwin Press, Inc.}} 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 {{convert|90|kg|abbr=on}} of sand each year. Or, on average (as there are so many variables i.e. size/species/location/depth etc.), almost {{convert|250|g|abbr=on|0}} per parrotfish per day.

While feeding, parrotfish must be cognizant of predation by one of their main predators, the lemon shark.{{cite book|last=Bright|first=Michael|title=The private life of sharks : the truth behind the myth|year=2000|publisher=Stackpole Books|location=Mechanicsburg, PA|isbn=978-0-8117-2875-1}} On Caribbean coral reefs, parrotfish are important consumers of sponges.{{Cite journal|title = Video-monitored predation by Caribbean reef fishes on an array of mangrove and reef sponges|journal = Marine Biology|date = 1996|pages = 117–123|volume = 126|doi = 10.1007/BF00571383|first1 = M|last1 = Dunlap|first2 = JR|last2 = Pawlik|s2cid = 84799900}} 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.{{Cite journal|title = Chemical defenses and resource trade-offs structure sponge communities on Caribbean coral reefs|journal = Proceedings of the National Academy of Sciences|date = 2014|pmid = 24567392|pages = 4151–4156|volume = 111|issue = 11|doi = 10.1073/pnas.1321626111|first1 = T-L|last1 = Loh|first2 = JR|last2 = Pawlik |pmc=3964098|bibcode = 2014PNAS..111.4151L|doi-access = free}}{{Cite journal|title = Indirect effects of overfishing on Caribbean reefs: sponges overgrow reef-building corals|last = Loh|first = TL|date = 2015|journal = PeerJ|doi = 10.7717/peerj.901|display-authors=etal|volume=3|pages=e901|pmid=25945305|pmc=4419544 | doi-access=free }}

Analysis of parrotfish feeding biology describes three functional groups: excavators, scrapers and browsers. Excavators have larger, stronger jaws that can gouge the substrate,{{Cite journal|last1=Price|first1=Samantha A.|last2=Wainwright|first2=Peter C.|last3=Bellwood|first3=David R.|last4=Kazancioglu|first4=Erem|last5=Collar|first5=David C.|last6=Near|first6=Thomas J.|date=2010-10-01|title=Functional Innovations and Morphological Diversification in Parrotfish|journal=Evolution|volume=64|issue=10|pages=3057–3068|doi=10.1111/j.1558-5646.2010.01036.x|pmid=20497217|s2cid=19070148|issn=1558-5646}} 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 epiphytes. 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 Scarus, 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.Environmental Biology of Fishes 28: 189–214, 1990

Recently, the microphage feeding hypothesis challenged the prevailing paradigm of parrotfish as algal consumers by proposing that:

{{Blockquote|text=Most parrotfishes are microphages that target cyanobacteria and other protein-rich autotrophic microorganisms that live on (epilithic) or within (endolithic) calcareous substrata, are epiphytic on algae or seagrasses, or endosymbiotic within sessile invertebrates.{{Cite journal|last1=Clements|first1=Kendall D.|last2=German|first2=Donovan P.|last3=Piché|first3=Jacinthe|last4=Tribollet|first4=Aline|last5=Choat|first5=John Howard|date=November 2016|title=Integrating ecological roles and trophic diversification on coral reefs: multiple lines of evidence identify parrotfishes as microphages|journal=Biological Journal of the Linnean Society|doi=10.1111/bij.12914}}}}

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.Georgina M Nicholson, Kendall D Clements, Micro-photoautotroph predation as a driver for trophic niche specialization in 12 syntopic Indo-Pacific parrotfish species, Biological Journal of the Linnean Society, Volume 139, Issue 2, June 2023, Pages 91–114, https://doi.org/10.1093/biolinnean/blad005 Additional microscopy and molecular barcoding research indicates that some parrotfish may ingest microscopic biota associated with endolithic sponges.Nicholson, G.M., Clements, K.D. A role for encrusting, endolithic sponges in the feeding of the parrotfish Scarus rubroviolaceus? Evidence of further trophic diversification in Indo-Pacific Scarini. Coral Reefs (2024). https://doi.org/10.1007/s00338-024-02482-z

Life cycle

File:Papageifische im Roten Meer..DSCF0262BE.jpg (Scarus ferrugineus) fighting.]]

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.{{Citation needed|date=January 2025}}

As pelagic spawners, parrotfish release many tiny, buoyant eggs into the water, which become part of the plankton. The eggs float freely, settling into the coral until hatching.{{Citation needed|date=January 2025}}

= Sex change =

File:Cetoscarus bicolor by Jacek Madejski.jpg (Cetoscarus bicolor) was described by Eduard Rüppell in 1829. In 1835, he mistakenly described the terminal phase, featured on this photo, as a separate species, C. pulchellus]]

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. [http://www.flmnh.ufl.edu/fish/Gallery/Descript/Sparrotfish/SParrotfish.html Stoplight parrotfish.] {{Webarchive|url=https://web.archive.org/web/20160120170504/https://www.flmnh.ufl.edu/fish/Gallery/Descript/SParrotfish/SParrotfish.html |date=20 January 2016 }} 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).{{cite journal | last1 = de Girolamo | first1 = Scaggiante | last2 = Rasotto | year = 1999 | title = Social organization and sexual pattern in the Mediterranean parrotfish Sparisoma cretense (Teleostei: Scaridae) | journal = Marine Biology | volume = 135 | issue = 2| pages = 353–360 | doi = 10.1007/s002270050634 | s2cid = 85428235 }}{{cite journal | last1 = Sadovy | last2 = Shapiro | year = 1987 | title = Criteria for the diagnosis of hermaphroditism in fishes | journal = Copeia | volume = 1987 | issue = 1| pages = 136–156 | doi = 10.2307/1446046 | jstor = 1446046 }} 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.{{cite book | author=Debelius, H. | year=1997 | title=Mediterranean and Atlantic Fish Guide: From Spain to Turkey - From Norway to South Africa | publisher=ConchBooks | page=221 | isbn=978-3925919541 }} In most species, juveniles have a different color pattern from adults. Juveniles of some tropical species can alter their color temporarily to mimic other species.Cardwell JR1, Liley NR.Gen Comp Endocrinol. 1991 Jan;81(1):7-20 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.

{{multiple image

| total_width = 440

| image1 = Scarus psittacus, Kahaluu-Keauhou, HI 96740, USA imported from iNaturalist photo 301406558 (cropped).jpg

| caption1 = Female Scarus psittacus (= initial phase)

| image2 = Scarus psittacus, Island of Hawai'i, Hawaii, USA imported from iNaturalist photo 63881409 (cropped).jpg

| caption2 = Male Scarus psittacus (= terminal phase)

}}

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.{{citation needed|date=March 2018}}

Economic importance

A commercial fishery exists for some of the larger species, particularly in the Indo-Pacific, but also for a few others like the Mediterranean parrotfish.{{cite journal | author=Cardigos, F. | year=2001 | title=Vejas | url=http://www.horta.uac.pt/projectos/marov/Media/Vejas.pdf | journal=Revista Mundo Submerso | volume=58 | issue=V | pages=48–51 | archive-url=https://web.archive.org/web/20180708074412/http://www.horta.uac.pt/projectos/marov/Media/Vejas.pdf | archive-date=8 July 2018 |url-status=dead }} Protecting parrotfishes is proposed as a way of saving Caribbean coral reefs from being overgrown with seaweedMorelle, Rebecca (1 November 2007) [http://news.bbc.co.uk/1/hi/sci/tech/7069933.stm Parrotfish to aid reef repair]. BBC 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; it is the only one of thousands of reef fish species that regularly performs the task of scraping and cleaning inshore coral reefs.{{cite journal|author=Australian Geographic|url=http://www.australiangeographic.com.au/news/2014/09/single-keystone-species-may-be-key-to-reef-health|title=Single species may be key to reef health|date= September 2014}}

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

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id:paleocene value:rgb(0.99,0.65,0.37)

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from: -65.5 till: -55.8 color:paleocene text:Paleocene

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

from: -23.03 till: -5.332 color:miocene text:Miocene

from: -5.332 till: -2.588 color:pliocene text:Plio.

from: -2.588 till: -0.0117 color:pleistocene text:Pleist.

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from: -23.03 till: -5.332 color:miocene text:Miocene

from: -5.332 till: -2.588 color:pliocene text:Plio.

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References

{{Reflist|35em|refs=

{{cite journal|title=Spatial patterns in reproductive traits of the temperate parrotfish Sparisoma cretense|url=http://www.horta.uac.pt/ppl/tmorato/pdf/Afonso_etal_2008a.pdf|doi=10.1016/j.fishres.2007.09.029|year=2008|last1=Afonso|first1=Pedro|last2=Morato|first2=Telmo|last3=Santos|first3=Ricardo Serrão|journal=Fisheries Research|volume=90|issue=1–3|pages=92–99}}

{{cite journal

|last=Videlier |first=H.

|author2=Geertjes, G.J. |author3= Videlier, J.J.

|year=1999

|title=Biochemical characteristics and antibiotic properties of the mucous envelope of the queen parrotfish

|journal=Journal of Fish Biology

|volume=54

|pages=1124–1127

|doi=10.1111/j.1095-8649.1999.tb00864.x

|issue=5}}

{{cite book

|editor=Paxton, J.R. |editor2=Eschmeyer, W.N.

|author1=Choat, J.H. |author2=Bellwood, D.R.

|name-list-style=amp |year=1998

|title=Encyclopedia of Fishes

|publisher= Academic Press

|location=San Diego

|pages= 209–211

|isbn= 978-0-12-547665-2}}

{{cite journal|author=Streelman, J. T., Alfaro, M. E.|display-authors=etal|year=2002|title=Evolutionary History of The Parrotfishes: Biogeography, Ecomorphology, and Comparative Diversity|journal=Evolution|volume=56|issue=5|pages=961–971|pmid=12093031|doi=10.1111/j.0014-3820.2002.tb01408.x|s2cid=41840374|df=dmy-all|doi-access=free}}

{{cite journal|author=Bellwood, D. R., Hoey, A. S., Choat, J. H. |year=2003|title=Limited functional redundancy in high diversity systems: resilience and ecosystem function on coral reefs|journal= Ecology Letters |volume=6|issue=4|pages= 281–285|doi=10.1046/j.1461-0248.2003.00432.x}}

{{cite journal|author=Lokrantz, J., Nyström, Thyresson, M., M., C. Johansson|title=The non-linear relationship between body size and function in parrotfishes|doi=10.1007/s00338-008-0394-3|bibcode=2008CorRe..27..967L|year=2008|journal=Coral Reefs|volume=27|issue=4|pages=967–974|s2cid=37926874}}

Lieske, E., and Myers, R. (1999). Coral Reef Fishes. 2nd edition. Princeton University Press. {{ISBN|0-691-00481-1}}

{{cite book

|last1=Thurman |first1=H.V

|last2=Webber |first2=H.H.

|title=Marine Biology

|year=1984

|publisher=Charles E. Merrill Publishing

|pages=303–313

|chapter=Chapter 12, Benthos on the Continental Shelf

|chapter-url=http://www.geology.iupui.edu/academics/CLASSES/G130/reefs/MB.htm }} Accessed 2009-06-14.

{{cite journal|pmid=15955516|year=2005|last1=Westneat|first1=MW|last2=Alfaro|first2=ME|title=Phylogenetic relationships and evolutionary history of the reef fish family Labridae|volume=36|issue=2|pages=370–90|doi=10.1016/j.ympev.2005.02.001|journal=Molecular Phylogenetics & Evolution}}

}}

Further reading

  • Hoey and Bonaldo. [https://books.google.com/books?id=pVNPDwAAQBAJ The Biology of Parrotfishes]
  • Monod, Th., 1979. "Scaridae". pp. 444–445. In J.C. Hureau and Th. Monod (eds.) Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean (CLOFNAM). UNESCO, Paris. Vol. 1.
  • {{cite journal | last = Sepkoski | first = Jack | title = A compendium of fossil marine animal genera | journal = Bulletins of American Paleontology | volume = 363 | page = 560 | year = 2002 | url = https://www.biodiversitylibrary.org/page/10698872 | access-date = 2014-05-03}}
  • {{cite journal | last = Smith | first = J.L.B.| title = The parrotfishes of the family Callyodontidae of the Western Indian Ocean | journal = Ichthyological Bulletin, Department of Ichthyology, Rhodes University | volume = 1 | year = 1956 | hdl = 10962/d1018535}}
  • {{cite journal | last = Smith | first = J.L.B.| title = The identity of Scarus gibbus Ruppell, 1828 and of other parrotfishes of the family Callyodontidae from the Red Sea and the Western Indian Ocean | journal = Ichthyological Bulletin, Department of Ichthyology, Rhodes University | volume = 16 | year = 1959 | hdl = 10962/d1018777}}
  • Bullock, A.E. and T. Monod, 1997. "Myologie céphalique de deux poissons perroquets (Teleostei: Scaridae)". Cybium 21(2):173–199.
  • {{cite journal | last1 = Randall | first1 = John E.| last2 = Bruce| first2 = Robin W. | title = The parrotfishes of the subfamily Scarinae of the Western Indian Ocean with descriptions of three new species | journal = Ichthyological Bulletin|publisher=J.L.B. Smith Institute of Ichthyology, Rhodes University | volume = 47 | year = 1983 | hdl = 10962/d1019747}}