Editing Clownfish

{{Short description|Monotypic subfamily of fishes}}
{{use dmy dates|date=February 2023}}
{{Automatic_taxobox
| image = Amphiprion ocellaris (Clown anemonefish) by Nick Hobgood.jpg
| image_caption = ''[[Amphiprion ocellaris]]''
| taxon = Amphiprion
| display_parents = 3
| authority = [[Marcus Elieser Bloch|Bloch]] & [[Johann Gottlob Schneider|Schneider]], 1801
| parent_authority = [[Gerald R. Allen|Allen]], 1975
| synonyms = *''Premnas'' <small>[[Georges Cuvier|Cuvier]], 1816</small>
}}
'''Clownfishes''' or '''anemonefishes''' are [[saltwater fish]]es in the genus '''''Amphiprion''''' within the subfamily '''Amphiprioninae'''. They are classified as damselfish within the family [[Pomacentridae]]. Clownfishes are found in the warm and tropical waters of the [[Indo-Pacific]], inhabiting mainly [[coral reef]]s. They have a distinctive colouration typically consisting of white vertical bars on a red, orange, yellow, brown or black background. Clownfish have developed a [[Mutualism (biology)|mutualistic]] [[Symbiosis|symbiotic]] relationship with [[sea anemone]]s, which they rely on for shelter and protection from predators. In turn, clownfish will protect the anemone from anemone-eating fish, as well as clean and fan them, and attract [[photosynthetic]] [[zooxanthellae]] via waste.  

Clownfish are [[omnivorous]] and mostly feed on [[plankton]]ic food. They live in groups consisting of a female and male breeding pair along with some non-breeding individuals. Clownfish have a size-based [[dominance hierarchy]] with the female ranking at the top, followed by the male and then the largest non-breeder and so on. When the female disappears, the male [[sequential hermaphroditism|changes sex]] and takes her place while the others move up the hierarchy. During reproduction, the female deposits the eggs on a rock near their anemone and the male [[External fertilization|fertilise]]s them. After hatching,clownfish disperse into the open ocean as larvae, and settle on the bottom and search for an anemone host as they become juveniles.  

The recognisable colour patterns and social nature of clownfish have contributed to their popularity. They are featured in the [[Disney/Pixar]] film ''[[Finding Nemo]]'' and are sought after in the [[Fishkeeping|aquarium trade]]. ''[[Amphiprion ocellaris]]'' is the most popular species and has been among the most commonly traded reef fish species. Many captive clownfish are taken from the wild and this has led to their decline. Clownfish are more numerous in [[marine protected area]]s, where collecting is forbidden. Other threats to populations include [[global warming]] which causes [[Ocean heat content|ocean warming]] and [[Ocean acidification|acidification]].

==Taxonomy==
Clownfish are damselfish (family [[Pomacentridae]]) in the genus ''Amphiprion'' (Greek {{lang|grc|amphi}}, 'on both sides' and {{lang|grc|prion}} 'saw'), which was coined by [[Marcus Elieser Bloch]] and [[Johann Gottlob Theaenus Schneider]] in 1801.<ref name=Tang2021/><ref>Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in {{harvnb|Frédérich|Parmentier|2016}} pp. 303, 306</ref> [[Georges Cuvier]] considered the [[maroon clownfish]] ([[specific name (zoology)|specific name]] ''biaculeatus'') morphologically different enough to be placed in its own genus ''Premnas'' in 1817.<ref>Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in {{harvnb|Frédérich|Parmentier|2016}} p. 306</ref> The status of ''Premnas'' has been disputed over the years, switching between a synonym or subgenus of ''Amphiprion'' and being its own genus. In 2021, two expansive [[phylogenetic]] analyses of damselfishes found the maroon clownfish to be within ''Amphiprion'', making ''Premnas'' a junior synonym.<ref name=Tang2021>{{cite journal |last1=Tang |first1=Kevin |last2=Stiassny |first2=Melanie |last3=Mayden |first3=Richard |last4=DeSalle |first4=Robert |title=Systematics of Damselfishes |journal=Ichthyology & Herpetology |date=May 2021 |volume=109 |issue=1 |page=258 |doi=10.1643/i2020105 |doi-access=free }}</ref><ref name=McCord2021/> In 1975, [[ichthyologist]] [[Gerald R. Allen]] placed clownfishes in their own subfamily Amphiprioninae.<ref>{{cite book|last=Allen|first=G. R.|year=1975|title=Damselfishes of the South Seas|publisher=TFH Publications|page=34|isbn=978-0876660348}}</ref> A 2009 study suggested creating the [[Tribe (biology)|tribe]] Amphiprionini for clownfishes and moving them to the subfamily [[Pomacentrinae]].<ref>{{cite journal|last1=Cooper|first1=W. J.|last2=Smith|first2=L. L.|last3=Westneat|first3=M. W.|year=2009|title=Exploring the radiation of a diverse reef fish family: phylogenetics of the damselfishes (Pomacentridae), with new classifications based on molecular analyses of all genera|journal=Molecular Phylogenetics and Evolution|volume=52|issue=1|pages=1–16|doi=10.1016/j.ympev.2008.12.010}}</ref> 

===Phylogeny===
The clownfish [[Lineage (evolution)|lineage]] diverged from other living damselfishes around 35 million years ago (mya) during the late [[Eocene]],<ref name=McCord2021/> and the [[most recent common ancestor]] is dated around 10.5 mya during the [[Early Miocene]]<ref name=Gaboriau2025/> in the waters of the [[Malay Archipelago]].<ref name=Litsios2014>{{cite journal|last1=Litsios|first1=G|last2=Pearman|first2=P. B.|last3=Lanterbecq|first3=D|last4=Tolou|first4=N|last5=Salamin|first5=N|year=2014|title=The radiation of the clownfishes has two geographical replicates|journal=Journal of Biogeography|volume=41|issue=11|pages=2140–2149|doi=10.1111/jbi.12370}}</ref> Clownfishes experienced an increase in species diversification starting around 5 mya,<ref name=McCord2021>{{cite journal|last1=McCord|first1=C. L.|last2=Nash|first2=C. M.|last3=Cooper|first3=W. J.|last4=Westneat|first4=M. W.|year=2021|title=Phylogeny of the damselfishes (Pomacentridae) and patterns of asymmetrical diversification in body size and feeding ecology|journal=PLoS ONE|volume=16|issue=10|page=e0258889|doi=10.1371/journal.pone.0258889|doi-access=free}}</ref> with two major [[adaptive radiation]]s; one centred on the Malay Archipelago and later in the waters of the western Indian Ocean.<ref name=Litsios2014/> High amounts of interbreeding between species occurred and the most recently diverged species are descended from hybrids.<ref name=hybrid>{{cite journal|last1=Litsios|first1=G|last2=Salamin|first2=N|year=2014|title=Hybridisation and diversification in the adaptive radiation of clownfishes|journal=BMC Evolutionary Biology|volume=14|page=245|doi=10.1186/s12862-014-0245-5|doi-access=free}}</ref> Clownfish speciation has been linked to their [[sea anemone]] hosts, species of which can be found in different habitats and thus drove ecological separation.<ref>{{cite journal|last1=Litsios|first1=G|last2=Sims|first2=C. A.|last3=Wüest|first3=R. O.|last4=Pearman|first4=P. B.|last5=Zimmermann|first5=N. E.|last6=Salamin|first6=N|year=2012|title=Mutualism with sea anemones triggered the adaptive radiation of clownfishes|journal=BMC Evolutionary Biology|volume=12|page=212|doi=10.1186/1471-2148-12-212|doi-access=free}}</ref><ref name=Gaboriau2025/>

There are 28 living species of clownfish and two hybrids forms, ''[[Amphiprion leucokranos]]'' (white-bonnet anemonefish) and ''[[Amphiprion thiellei]]'' (Thielle’s anemonefish).<ref name=preface>"Preface" in {{harvnb|Laudet|Ravasi|2023}} p. xii</ref> In 1972, Allen listed five major clades or [[species complex|complexes]] based on morphology; ''percula'',  ''akallopisos'', ''ephippium'', ''polymnus'' and ''clarkii'' with ''A. biaculeatus'' being in a [[Monotypic taxon|monotypic]] clade close to ''percula''.{{sfn|Allen|1975b|pp=46–52}}. A 2014 study lists two more major clades: Australian and Indian, with two species ''[[Amphiprion chrysopterus]]'' and ''[[Amphiprion latezonatus]]'' bring monotypic clades and ''A. biaculeatus'' under the ''percula'' clade.<ref name=hybrid/> A 2021 study placed ''A. chrysopterus'' and members of the clade ''polymnus'' in the Indian clade,<ref name=McCord2021/> while a 2025 study found the ''percula'' clade to be [[paraphyly|paraphyletic]].<ref name=Gaboriau2025/>

The following cladogram of the 28 clownfish species is based on a genetic study by Gaboriau and colleagues (2025):<ref name=Gaboriau2025>{{cite journal|last1=Gaboriau|first1=T|last2=Marcionetti|first2=A|last3=Garcia-Jimenez|first3=A|last4=Schmid|first4=S|last5=Fitzgerald|first5=L. M.|last6=Micheli|first6=B|last7=Titus|first7=B|last8=Salamin|first8=N|year=2025|title=Host use drives convergent evolution in clownfish|journal=Proceedings of the National Academy of Sciences|volume=122|issue=17|page=e2419716122|doi=10.1073/pnas.2419716122}}</ref>{{efn|Common names are taken from<ref name=preface/>}}

{{clade|style=font-size:85%; line-height:85%
|label1=''Amphiprion''
|1={{clade
 |1=''[[Maroon clownfish|A. biaculeatus]]'' (Maroon clownfish or Spinecheek anemonefish) [[File:Clownfish (Papua New Guinea).jpg|40px]]  
 |2={{clade
  |1={{clade
   |1=''[[Amphiprion percula|A. percula]]'' (Orange clownfish) [[File:Clown.fish.arp.750pix.jpg|40px]]
   |2=''[[Amphiprion ocellaris|A. ocellaris]]'' (False clownfish) [[File:Clown fish in the Andaman Coral Reef.jpg|40px]] }}
  |2={{clade
   |1=''[[Amphiprion latezonatus|A. latezoneatus]]'' (Wide-band anemonefish) [[File:Amphiprion latezonatus, Norfolk.jpg|40px]]
   |2={{clade
    |1={{clade
     |1={{clade
      |1=''[[Amphiprion perideraion|A. perideraion]]'' (Pink skunk anemonefish) [[File:Pink Anemonefish (8 cm).png|40px]]
      |2={{clade
       |1=''[[Amphiprion pacificus|A. pacificus]]'' (Pacific anemonefish) [[File:Amphiprion pacificus.webp|40px]]
       |2={{clade
        |1=''[[Amphiprion akallopisos|A. akallopisos]]'' (Skunk anemonefish) [[File:Amphiprion akallopisos 13376076.jpg|40px]]
        |2=''[[Amphiprion sandaracinos|A. sandaracinos]]'' (Yellow skunk anemonefish) [[File:Amphiprion sandaracinos Minatoyama2.jpg|40px]] }} }} }}
     |2={{clade
      |1={{clade
       |1=''[[Amphiprion tricinctus|A. tricinctus]]'' (Three-band anemonefish) [[File:Amphiprion tricinctus 11-2022.JPG|40px]]
       |2=''[[Clark's anemonefish|A. clarkii]]'' (Yellow-tail anemonefish) [[File:XRF-Amphiprion clarkii.png|40px]] }}
      |2={{clade
       |1=''[[Amphiprion barberi|A. barberi]]'' (Barber’s anemonefish) [[File:Amphiprion barberi.jpg|40px]]
       |2={{clade
        |1={{clade
         |1=''[[Amphiprion frenatus|A. frenatus]]'' (Tomato anemonefish) [[File:Tomato clownfish Amphiprion frenatus.jpg|40px]]
         |2=''[[Amphiprion ephippium|A. ephippium]]'' (Saddle anemonefish) [[File:Amphiprion ephippium (Bloch, 1790).jpg|40px]] }}
        |2={{clade
         |1=''[[Amphiprion melanopus|A. melanopus]]'' (Fire anemonefish) [[File:Amphiprion melanopus RLS.jpg|40px]]
         |2=''[[Amphiprion rubrocinctus|A. rubrocinctus]]'' (Red anemonefish) [[File:Amphiprion rubrocinctus RLS.jpg|40px]] }} }} }} }} }}
    |2={{clade
     |1={{clade
      |1=''[[Amphiprion chrysogaster|A. chrysopterus]]'' (Orange-fin anemonefish) [[File:Pomacentridae Amphiprion chrysogaster 1.jpg|40px]] 
      |2={{clade
       |1=''[[Amphiprion mccullochi|A. mccullochi]]'' (White-snout anemonefish) [[File:Amphiprion mccullochi RLS3.jpg|40px]]
       |2=''[[Amphiprion akindynos|A. akindynos]]'' (Barrier reef anemonefish) [[File:Amphiprion akindynos.jpg|40px]] }} }}
     |2={{clade
      |1={{clade
       |1=''[[Amphiprion sebae|A. sebae]]'' (Sebae anemonefish) [[File:XRF-Amphiprion sebae.png|40px]]
       |2=''[[Amphiprion polymnus|A. polymnus]]'' (Saddleback anemonefish) [[File:Amphiprion Species.JPG|40px]] }}
      |2={{clade
       |1={{clade
        |1=''[[Amphiprion nigripes|A. nigripes]]'' (Maldive anemonefish) [[File:XRF-Amphiprion nigripes.png|40px]]
        |2=''[[Amphiprion chagosensis|A. chagosensis]]'' (Chagos anemonefish) [[File:A, chagosensis 3.jpg|40px]] }}
       |2={{clade
        |1={{clade
         |1=''[[Amphiprion chrysogaster|A. chrysogaster]]'' (Mauritian anemonefish) [[File:Pomacentridae Amphiprion chrysogaster 1.jpg|40px]]
         |2=''[[Amphiprion fuscocaudatus|A. fuscocaudatus]]'' (Seychelles anemonefish) [[File:Amphiprion fuscocaudatus.jpg|40px]] }}
        |2={{clade
         |1={{clade
          |1=''[[Amphiprion bicinctus|A. bicinctus]]'' (Two-band anemonefish) [[File:Amphiprion bicinctus by Patryk Krzyzak.jpg|40px]]
          |2=''[[Amphiprion omanensis|A. omanensis]]'' (Oman anemonefish) [[File:Amphiprion omanensis.TIF|40px]] }}
         |2={{clade
          |1=''[[Amphiprion allardi|A. allardi]]'' (Two-bar anemonefish) [[File:Amphiprion allardi 221884898.jpg|40px]]
          |2=''[[Amphiprion latifasciatus|A. latifasciatus]]'' (Madagascar anemonefish) [[File:Clowns et oeufs à Mayotte.jpg|40px]] }} }} }} }} }} }} }} }} }} }} }}

==Characteristics==
Clownfish vary in size; ''A. biaculeatus'' can reach {{convert|160|mm|in|abbr=on}} long while ''[[Amphiprion percula|A. percula]]'' reaches only {{convert|80|mm|in|abbr=on}}.{{sfn|Fautin|Allen|1992|pp=98, 114}}. Females are larger than males and the smallest individuals in a group are only {{convert|6|mm|in|abbr=on}}.{{sfn|Allen|1975b|p=180}}<ref>Casas, L.; Parker, C. G.; Rhodes, J. S.; "Sex Change from Male to Female Active Feminization of the Brain, Behavior, and Gonads in Anemonefish" in {{harvnb|Laudet|Ravasi|2023}} p. 188</ref> Clownfishes vary from oval-shaped to streamlined, and have rounded heads that lack scales between the snout and eyes. Teeth are present on both the [[fish jaw|oral and pharyngeal jaws]] but absent on the [[palate]], and may be conical or chisel-shaped.{{sfn|Allen|1975b|pp=36, 184}} They have saw-shaped edges along the [[operculum (fish)|operculum]] (gill covering) and suborbital area, which is the source of their genus name. The [[dorsal fin]] has 10 spines and 14–20 soft [[Fish fin#Bony fishes|rays]];<ref>Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in {{harvnb|Frédérich|Parmentier|2016}} pp. 303</ref> ''[[Amphiprion ocellaris|A. ocellaris]]'' has a large recess between the spines and soft rays, while in ''[[red saddleback anemonefish|A. ephippium]]'' they are mostly continuous.<ref name=whitebar/> Rays number 15–21 in the [[pectoral fin]]s, five in the [[pelvic fin]]s, 11–15 in the [[anal fin]], and 14–15 in the [[caudal fin]].{{sfn|Allen|1975b|p=36}}

===Colour patterns===
Clownfish have distinctive colour patterns consisting of a red, orange, yellow, brown or black background with up to three white vertical bars lined with black. Some species have only a horizontal line along the back or both a horizontal and a vertical line; one species ''A. ephippium'', has no bars at all. Orange, yellow and red colouration is created by [[xanthophore]] [[chromatophore|pigment cells]], black and brown by [[melanophore]]s and the white bars by [[iridophore]]s. Vertical bar formation starts at the front: species with only one bar have it at the head, those with two at the head and trunk, and three at the head, trunk and tail. Variations in the number of vertical bars between individuals of the same species occurs in ''[[Amphiprion melanopus|A. melanopus]]'', ''[[Amphiprion polymnus|A. polymnus]]'' and ''[[Amphiprion clarkii|A. clarkii]]''.<ref name=whitebar>{{cite journal|last1=Salis|first1=P|last2=Roux|first2=N|last3=Soulat|first3=O|last4=Lecchini|first4=D|last5=Laudet|first5=V|last6=Frédérich|first6=B|year=2018|title=Ontogenetic and phylogenetic simplification during white stripe evolution in clownfishes|journal=BMC Biology|volume=16|page=90|doi=10.1186/s12915-018-0559-7|doi-access=free}}</ref><ref name=Salis65>Salis, P.; Klann, M.; Laudet, V. "Color Pattern in Anemonefish: Develpment, Role and Diversity" in {{harvnb|Laudet|Ravasi|2023}} pp. 65–67</ref> Numerous [[Polymorphism (biology)|color morph]] mutations occur particularly in captive clownfish, including [[melanism]], a "Misbar" (incomplete vertical bars) morph, and a "golden" morph which is caused by a lack of both melanophores and iridophores. There are also morphs with thickened and merged bars.<ref>Salis, P.; Klann, M.; Laudet, V. "Color Pattern in Anemonefish: Develpment, Role and Diversity" in {{harvnb|Laudet|Ravasi|2023}} pp. 72–73</ref>

Merilata and colleagues (2018) found that clownfish with only one or no vertical bars tend to be more [[Generalist and specialist species|specialised]] for anemone species with greater toxicity and shorter tentacles. Conversely, clownfish species with two or three bars are more likely to use more anemone species within their range, several of which have longer tentacles. The researchers suggest that vertical bars function in [[camouflage]] while [[aposematism|warning colouration]] is more important for species that cannot hide in the tentacles of their hosts. This would be a unique case of aposematism since it warns about another animal, namely the anemone.<ref name=Merilata2018>{{cite journal|last1=Merilata|first1=S|last2=Kelley|first2=J. L.|year=2018|title= Scary clowns: adaptive function of anemonefish coloration|journal=Journal of Evolutionary Biology|volume=31|issue=10|pages=1558–1571|doi=10.1111/jeb.13350}}</ref> Salis and colleagues (2018) suggested that the bars function in [[Intra-species recognition|species recognition]], finding little [[sympatry|overlap]] between species with the same bar numbers within various clownfish [[Community (ecology)|communities]].<ref name=whitebar/> A 2024 study found evidence for this function as ''A. ocellaris'' can distinguish  between individuals of different bar numbers.<ref name=Hayashi2024/>  Merilata and colleagues (2018) dispute this, noting the geographic and ecological overlap between the similar-looking ''A. percula'' and ''A. ocellaris''.<ref name=Merilata2018/> 

<gallery mode="packed" heights="150px">
File:Amphiprion ocellaris (Clown anemonefish) Nemo.jpg|''[[Amphiprion ocellaris|A. ocellaris]]'' with three vertical bars
File:Diving at Siaes Tunnel, Palau.jpg|''[[Amphiprion allardi|A. allardi]] '' with two vertical bars
File:Amphiprion mccullochi pareja.jpg|''[[Amphiprion mccullochi|A. mccullochi]]'' with one vertical bar
File:Amphiprion ephippium.jpg|''[[Amphiprion ephippium|A. ephippium]]'' with no bars
File:Pink Anemonefish - Bunaken.jpg|''[[Amphiprion perideraion|A. perideraion]]'' with one vertical bar and one horizontal dorsal bar
File:Amphiprion sandaracinos.jpg|''[[Amphiprion sandaracinos|A. sandaracinoss]]'' with a horizontal dorsal bar
</gallery>

==Distribution and habitat==
Clownfishes inhabit warm and tropical waters spanning the Indian Ocean and the western Pacific; from the [[Red Sea]] to [[French Polynesia]], and from Japan down to Australia.<ref name=Colleye304/><ref>Hayashi, K.; Riemer, J. D. "Habitat Selection of Anemonefish" in {{harvnb|Laudet|Ravasi|2023}} p. 169</ref> Some species are more widespread than others, and some live only around islands or [[archipelago]]s. Areas closer to the edges of their distribution have fewer species; both the Red Sea and French Polynesia have only one species each. By contrast, as many as six species inhabit the waters of [[Great Barrier Reef]].<ref name=Colleye304>Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in {{harvnb|Frédérich|Parmentier|2016}} pp. 304–305</ref> Clownfishes are limited by the distribution of their sea anemone hosts. Hence they are found mainly at the [[photic zone]]s where there is more [[photosynthetic]] [[zooxanthellae]] for their hosts to depend on. This includes [[coral reefs]] and surrounding areas.<ref name=Fautin1991>{{cite journal|last1=Fautin|first1=D. G.|year=1991|title=The anemonefish symbiosis: what is known and what is not|journal=Symbiosis|volume=10|pages=23–46|url=https://kuscholarworks.ku.edu/entities/publication/ed3447a6-e5c6-4e01-a398-d22b5cbe3b19}}</ref> Within an reef, clownfish species that use the same anemone species as their main hosts will avoid competition by using individuals at different zones (nearshore, mid-[[lagoon]], and outer bank).<ref>{{cite journal|last1=Elliott|first1=J. K.|last2=Mariscal|first2=R. N.|year=2001|title=Coexistence of nine anemonefish species: differential host and habitat utilization, size and recruitment|journal=Marine Biology|volume=138|pages=23–36|doi=10.1007/s002270000441}}</ref>

==Behaviour and ecology==
[[File:Clownfish in anemone off Vanuatu.ogg|thumb|right|Video of a [[cinnamon clownfish|A. melanopus]] swimming around an anemone]]
===Feeding===
Clownfish are [[omnivorous]], and mostly feed on [[plankton]]ic food such as [[copepod]]s and larval [[tunicate]]s. [[Algae]] is also an important food source and make up much of the diet of ''A. perideraion''. Clownfish will also feed on the waste dischanged by the anemone.{{sfn|Allen|1975b|pp=180, 184–189}}{{sfn|Fautin|Allen|1992|p=132}} Feeding takes up most of a clownfish's acivity; around 90 percent in dominant ''A. chrysopterus'' specifically. Where predators are less common, clownfish may forage as far as {{convert|20|m2|abbr=on}} of ocean floor away from their anemone. Otherwise they are restricted to feeding in the water column above their host.{{sfn|Allen|1975b|p=202}} The dominant pair in a clownfish group feed further from the anemone than the smaller subordinates.{{sfn|Fautin|Allen|1992|p=132}}

===Mutualism===
{{further information|Symbiosis|Mutualism (biology)}}
Clownfish have a [[Mutualism (biology)|mutualistic]] [[Symbiosis|symbiotic]] relationship with sea anemones.{{sfn|Allen|1975b|p=180}}<ref name=Colleye312>Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in {{harvnb|Frédérich|Parmentier|2016}} p. 312–313</ref> They acclimate themselves to their hosts by touching, nipping and fanning the tentacles over a period of minutes to days.<ref name=Hoepner201/> The main benefit of living among anemones is protection from predators by anemone's stinging tentacles. Straying clownfish retreat to the safety of the tentacles when they encounter a potential threat and they are always near their hosts, with smaller fish rarely leaving the [[Sea anemone#Anatomy|oral disc]].<ref name=Fautin1991/>{{sfn|Allen|1975b|p=180}} Clownfish may even swim into the [[coelenteron]] (gastrovascular cavity), though Gerald Allen observes this to be uncommon. Nighttime is spend resting deep among the tentacles.{{sfn|Allen|1975b|p=180}} A less important benefit for clownfish is nourishment from the discharged waste and parasites.<ref name=Fautin1991/>{{sfn|Allen|1975b|p=180}} 
[[File:Clark's Anemonefish (6997549061).jpg|thumb|right|''[[Amphiprion clarkii|A. clarkii]]'' with an ''[[Entacmaea quadricolor|E. quadricolor]]'' anemone, the species utilizes all ten anemone species which host clownfish]]
Anemones are less dependent on clownfish than the fish are of them; as evident as many individuals of host species lack clownfish.<ref name=Fautin1991/> Nevertheless, clownfish contribute to the survival of their hosts by guarding from anemone-eating fish such as the [[butterflyfish]] species ''[[Chaetodon lunula]]''.<ref name=Fautin1991/>{{sfn|Allen|1975b|p=183}} Other benefits they provide include the cleaning off of copepod parasites, increased oxygen flow via the rapid movements of the fish's fins and the attraction of more zooxanthellae by clownfish waste.<ref name=Colleye312/><ref>{{cite journal|last1=Szczebak|first1=J. T.|last2=Henry|first2=R. P.|last3=Al-Horani|first3=F. A.|last4=Chadwick|first4=N. E.|year=2013|title=Anemonefish oxygenate their anemone hosts at night|journal=Journal of Experimental Biology|volume=216|issue=6|pages=970–976|doi=10.1242/jeb.075648}}</ref> A 2005 study found that anemone grew and regenerated faster with clownfish them without, and attributed this to [[ammonium]] from clownfish waste.<ref name="D. Porat">{{cite journal |last1=Porat |first1=D. |last2=Chadwick-Furman |first2=N.E. |title=Effects of anemonefish on giant sea anemones: Ammonium uptake, zooxanthella content and tissue regeneration |journal=Marine and Freshwater Behaviour and Physiology |date=March 2005 |volume=38 |issue=1 |pages=43–51 |doi=10.1080/10236240500057929 |bibcode=2005MFBP...38...43P |s2cid=53051081 }}</ref>

A total of ten sea anemone species are used by clownfish as hosts: ''[[Radianthus malu]]'', ''[[Radianthus crispa|R. crispa]]'', ''[[Radianthus magnifica|R. magnifica]]'', ''[[Stichodactyla mertensii]]'', ''[[Stichodactyla haddoni|S. haddoni]]'', ''[[Stichodactyla gigantea|S. gigantea]]', [[Cryptodendrum adhaesivum]]'', ''[[Entacmaea quadricolor]]'', ''[[Heteractis aurora]]'' and ''[[Macrodactyla doreensis]]''. Some clownfish are [[Generalist and specialist species|generalist]] in their choice of hosts while others are more specialised. ''[[Clark's anemonefish|A. clarkii]]'' is the most generalised species and utilises all ten anemone species, while nine — ''[[Amphiprion frenatus|A. frenatus]]'', ''[[Amphiprion chagosensis|A. chagosensis]]'', ''[[Amphiprion pacificus|A. pacificus]]'', ''[[Amphiprion fuscocaudatus|A. fuscocaudatus]]'', ''[[Amphiprion latifasciatus|A. latifasciatus]]'', ''[[Amphiprion mccullochi|A. mccullochi]]'', ''[[Amphiprion nigripes|A. nigripes]]'', ''[[Amphiprion sebae|A. sebae]]'', and ''A. biaculeatus'' — use just one anemone species. Desirable traits in a host include long tentacles to hide among. In addition, certain anemones like ''H. aurora'' and ''E. quadricolor'' have tentacles with knob-like structures which provide more surface area for the fish to conceal itself. ''R. magnifica'' can provide extra protection when it pulls all its tentacles inside a soft body. The potency of venom is also a desirable trait; highly toxic anemone species tend to have smaller tentacles and so provide less shelter but more protection.<ref name=Hoepner198>Hoepner, C. M.; Fobert, E. K.; Abbott, C. A.; da Silva, K. B. "No Place Like Home: Can Omics Uncover the Secret behind the Sea Anemone and Anemonefish Symbiotic Relationship?" in {{harvnb|Laudet|Ravasi|2023}} pp. 198–200</ref>

Their ability to avoid being stung is attributed to their mucus coating.<ref name=Fautin1991/> There is evidence that clownfish mucus mimics the molecules or bacteria of anemone mucus and lacks  trigger for the anemone’s [[nematocyst]]s (stinging barbs). Mucus thickness may also play a role, but this is not clear.<ref name=Hoepner201>Hoepner, C. M.; Fobert, E. K.; Abbott, C. A.; da Silva, K. B. "No Place Like Home: Can Omics Uncover the Secret behind the Sea Anemone and Anemonefish Symbiotic Relationship? in {{harvnb|Laudet|Ravasi|2023}} pp. 201–204</ref> There is dispute over how much of the mucus is innate to the clownfish and how much is gained from the anemone during the acclimation period.<ref name=Fautin1991/> Nguyen and colleagues (2023) write "Whereas some anemone fish species seem to produce their own protective mucous coating, others may acquire mucus (or biomolecules within) from the sea anemone during an acclimation period".<ref>{{cite journal|last1=Nguyen|first1=H-T. T|last2=Zhao|first2=M|last3=Wang|first3=T|last4=Dang|first4=B. T.|last5=Geffen|first5=A. J.|last6=Cummins|first6=S. F.|year=2023|title=Sea anemone–anemone fish symbiosis: behavior and mucous protein profiling|journal=Journal of Fish Biology|volume=105|issue=2|pages=603–618|doi=10.1111/jfb.15772}}</ref> Roux and colleagues (2019) found evidence that clownfish exchange [[microbiota]] with their anemone hosts.<ref>{{cite journal|last1=Roux|first1=N|last2=Lami|first2=R|last3=Salis|first3=P|last4=Magré|first4=K|last5=Romans|first5=P|last6=Masanet|first6=P|last7=Lecchini|first7=D|last8=Laudet|first8=V|year=2019|title=Sea anemone and clownfish microbiota diversity and variation during the initial steps of symbiosis|journal=Scientific Reports|volume=9|page=19491|doi=10.1038/s41598-019-55756-w}}</ref>

===Social structure===
[[File:Anemonefish komodo.jpg|thumb|right|Group of ''[[Amphiprion perideraion|A.  perideraion]]'', including the dominant pair and non-breeding individuals]]
A group of clownfish occupying an anemone usually consists of a breeding female and male along with some non-breeding individuals.<ref name=Beldade130/> [[Dominance hierarchy|Dominance]] in clownfish groups is based on size, with the female being the largest and most dominant, followed by the male and then the largest non-breeder and so on. An individual is 20 percent larger than its direct subordinate.<ref name=Beldade130/><ref name=Buston2004/> In ''A. percula'', the number of non-breeders ranges from zero to four, with group size depending on anemone size,<ref name=Buston2003>{{cite journal|last1=Buston|first1=P|year=2003|title=Forcible eviction and prevention of recruitment in the clown anemonefish|journal=Behavioral Ecology|volume=14|issue=4|pages=576–582|doi=10.1093/beheco/arg036}}</ref> as well as the size of the female as larger females allow for more members without unbalancing the size ratio between them.<ref>{{cite journal|last1=Buston|first1=P. M.|last2=Cant|first2=M. A.|year=2006|title=A new perspective on size hierarchies in nature: patterns, causes, and consequences|journal=Oecologia|volume=149|issue=2|pages=362–372|doi=10.1007/s00442-006-0442-z}}</ref> Members of a group are unrelated.<ref>{{cite journal|last1=Buston|first1=P. M.|last2=Bogdanowicz|first2=S. M.|last3=Wong|first3=A|last4=Harrison|first4=R. G.|year=2007|title=Are clownfish groups composed of close relatives? An analysis of microsatellite DNA variation in ''Amphiprion percula''|journal=Molecular Ecology|volume=16|issue=17|pages=3671–3678|doi=10.1111/j.1365-294X.2007.03421.x}}</ref>

As [[Sequential hermaphroditism|protandrous sequential hermaphrodite]]s, the male clownfish changes into a female when the previous one is lost, while the largest non-breeder becomes a male and the others rise in rank.<ref name=Beldade130>Beldade, R.; Bernardi, G.; Mills, S. C. "Anemonefish Behavior and Reproduction" in {{harvnb|Laudet|Ravasi|2023}} pp. 130</ref><ref name=Buston2004/> New fish that join the group rank at the bottom.<ref name=Buston2004>{{cite journal|last1=Buston|first1=P. M.|year=2004|title=Territory inheritance in clownfish|journal=Proceedings of the Royal Society B: Biological Science|volume=271|pages=S252–S254|doi=10.1098/rsbl.2003.0156}}</ref> Non-breeders are forced to wait for their time to become breeders, since nearby anemones are occupied and they are too small to challenge the dominants.<ref>{{cite journal|last1=Branconi|first1=R|last2=Barbasch|first2=T. A.|last3=Francis|first3=R. K.|last4=Srinivasan|first4=M|last5=Jones|first5=G. P.|last6=Buston|first6=P. M.|year=2020|title=Ecological and social constraints combine to promote evolution of non-breeding strategies in clownfish|journal=Communications Biology|volume=3|page=649|doi=10.1038/s42003-020-01380-8}}</ref> The dominant pair controls membership of the group and will drive away individuals when the anemone gets too full,<ref name=Buston2003/> particularly those that are close to them in size. Thus newcomers must remain smaller than their immediate superior to avoid getting evicted.<ref name=Buston2004/> Clownfish maintain their dominance hierarchy via displays, sound production and chasing. Sounds produced by clownfish include "clicks", "grunts", "pops" and "chirps". Dominants will chase their subordinates while producing a sound consisting of one or more long pulses. The subordinate submits to by emitting a sound with quicker pulses while shaking their heads.<ref>{{cite journal|last1=Colleye|first1=O|last2=Parmentier|first2=E|year=2012|title=Overview on the diversity of sounds produced by clownfishes (Pomacentridae): importance of acoustic signals in their peculiar way of life|journal=PLoS ONE|volume=7|issue=11|page=e49179|doi=10.1371/journal.pone.0049179|doi-access=free}}</ref> Clownfish appear to produce sounds via the jaws, which is amplified by the swim bladder.<ref>Parmentier, E.; Lecchini, D. "Sound Communication" in {{harvnb|Laudet|Ravasi|2023}} p. 100</ref><ref>{{cite journal|last1=Parmentier|first1=E|last2=Colleye|first2=O|last3=Fine|first3=M. L.|last4=Frédérich|first4=B|last5=Vandewalle|first5=P|last6=Herrel|first6=A|year=2007|title=Sound production in the clownfish ''Amphiprion clarkii''|journal=Science|volume=316|issue=5827|page=1006|doi=10.1126/science.1139753}}</ref>

One study of captive ''A. ocellaris'' found that the dominant pair are the most territorial while non-breeders are much less so. Both the male and female direct their aggression against intruders of the same sex, though resident males are more likely display than attack. Similarly non-breeding intruders are more likely to be simply intimidated.<ref>{{cite journal|last1=Iwata|first1=E|last2=Manbo|first2=J|year=2013|title=Territorial behaviour reflects sexual status in groups of false clown anemonefish (''Amphiprion ocellaris'') under laboratory conditions|journal=acta ethologica|volume=16|pages=97–103|doi=10.1007/s10211-012-0142-0}}</ref> Another study of the same species found they showed more aggression towards fish that have three vertical bars followed by those with two, then one and none, suggesting that it recognises and sees members of their species as their main competition for anemones.<ref name=Hayashi2024>{{cite journal|last1=Hayashi|first1=K|last2=Locke|first2=N. J. M.|last3=Laudet|first3=V|year=2024|title=Counting Nemo: anemonefish ''Amphiprion ocellaris'' identify species by number of white bars|journal=Journal of Experimental Biology|volume=227|issue=2|page=jeb246357|doi=10.1242/jeb.246357}}</ref> Conversely, ''A clarkii'' was observed to attack individuals of other species more than those of its own.<ref>{{cite journal|last=Hattori|first=A|year=2022|title=Small and large anemonefishes can coexist using the same patchy resources on a coral reef, before habitat destruction|journal=Journal of Animal Ecology|volume=71|issue=5|pages=824–831|doi=10.1046/j.1365-2656.2002.00649.x}}</ref>

<gallery mode="packed" heights="150px">
File:Amphiprion frenatus aggressive sounds - journal.pone.0049179.s001.ogv|''A. clarkii'' dominant chasing subordinate while producing aggressive sounds
File:Amphiprion frenatus submissive sounds - journal.pone.0049179.s002.ogv|''A. frenatus'' subordinate head-shaking while producing submissive sounds
</gallery>

===Reproduction and lifecycle===
Clownfish breed year-round in tropical waters while in more temperate waters, like those around Japan, breeding occurs mostly in spring and summer. Only the dominant female and male reproduce which mostly occurs during a [[full moon]]. In the days leading up to spawning, the pair perform courtship rituals which involve the male chasing and nibbling the female as well as erecting his dorsal pelvic and anal fins while staying motionless in front or alongside her. Both the female and male then prepare a nest by cleaning up a nearby rock. Here the female will deposit eggs for the male to [[External fertilization|fertilise]]. Clownfish lay up to a thousand eggs, which are conical in shape, {{convert|3|–|4|mm|abbr=on}} long and stick to the rocky substrate by bundles of short fibres.{{sfn|Fautin|Allen|1992|pp=126–129}}<ref name=Beldade132>Beldade, R.; Bernardi, G.; Mills, S. C. "Anemonefish Behavior and Reproduction" in {{harvnb|Laudet|Ravasi|2023}} p. 132</ref> The male tends to the fertilised eggs, cleaning and guarding them as well as fanning them with his pectoral fins.{{sfn|Fautin|Allen|1992|p=129}} 

Incubation lasts six to seven days.{{sfn|Fautin|Allen|1992|p=129}} The eggs start out bright orange and progressively darken, and the eyes of the embryos develop and become visible. The fish break out of their capsules during nighttime.<ref name=Beldade132/> After hatching, clownfish enter the larval and [[pelagic]] stage of their development. This stage lasts up to 12 days which is shorter than that of other damselfishes can last for 70 days.{{sfn|Fautin|Allen|1992|p=130}}<ref name=Roux2019>{{cite journal|last1=Roux|first1=N|last2=Salis|first2=P|last3=Lambert|first3=A|last4=Logeux|first4=V|last5=Soulat|first5=O|last6=Romans|first6=P|last7=Frédérich|first7=B|last8=Lecchini|first8=D|last9=Laudet|first9=V|year=2019|title=Staging and normal table of postembryonic development of the clownfish (''Amphiprion ocellaris'')|journal=Development Dynamtics|volume=248|issue=7|pages=545–568|doi=10.1002/dvdy.46}}</ref> Larval clownfish are initially transparent, expect for the eyes, yolk-sac and some pigment spots.{{sfn|Fautin|Allen|1992|p=130}} Over time they begin to [[metamorphosis|metamorphosise]]; growing in size and developing their fins, sensory and internal organs, [[notochord]] flexion and colouration.<ref name=Roux2019/> Clownfish larvae can [[biological dispersal|disperse]] widely across open ocean; ''[[Amphiprion omanensis|A. omanesis]]'' has been recorded travelling over {{convert|400|km|abbr=on}} along ocean currents.<ref>{{cite journal|last1=Simpson|first1=S. D.|last2=Harrison|first2=S. D.|last3=Claereboudt|first3=M. R.|last4=Planes|first4=S|year=2014|title=Long-distance dispersal via ocean currents connects Omani clownfish populations throughout entire species range|journal=PLoS ONE|volume=9|issue=9|page=e107610|doi=10.1371/journal.pone.0107610|doi-access=free}}</ref> 

As they enter the juvenile stage, clownfish begin settle to the ocean floor and find an anemone host,<ref name=Roux2019/> while transitioning to a more [[Diurnality|diurnal]] lifestyle.<ref>{{cite journal|last1=Schalm|first1=G|last2=Bruns|first2=K|last3=Drachenberg|first3=N|last4=Geyer|first4=N|last5=Foulkes|first5=N. S.|last6=Bertolucci|first6=C|last7=Gerlach|first7=G|year=2021|title=Finding Nemo’s clock reveals switch from nocturnal to diurnal activity|journal=Scientific Reports|volume=11|page=6801|doi=10.1038/s41598-021-86244-9}}</ref> Juveniles continue to grow and develop their adult colouration,<ref name=Roux2019/><ref name=whitebar/> but cannot produce [[gamete]]s until they ascend to dominance within a group.<ref name=Roux2019/> The dominant male's gonads produce sperm but also possess dormant ovarian cells. When transitioning into a female, the gonads switch to producing ovaries.{{sfn|Fautin|Allen|1992|p=131}} The transition from male to female starts with an increases in body size and feminisation of the brain, followed by gonadal changes and then behaviour changes. The process can last over four months.<ref>Casas, L.; Parker, C. G.; Rhodes, J. S. "Sex Change from Male to Female: Active Feminization of the Brain, Behavior, and Gonads in Anemonefish" in {{harvnb|Laudet|Ravasi|2023}} p. 119</ref> Clownfish can live for over 20 years.<ref>Mutalipassi, M.; Tozzini, E. T.; Cellerino, A. "Age and Longevity" in {{harvnb|Laudet|Ravasi|2023}} pp. 79–80</ref> ''A. percula'' is estimated to reach 30 years; twice as much as the average reef damselfish and six times that of a fish its size.<ref>{{cite journal|last1=Buston|first1=P. M.|last2=Garcia|first2=M. B.|year=2007|title=An extraordinary life span estimate for the clown anemonefish ''Amphiprion percula''|journal=Journal of Fish Biology|volume=70|issue=6|pages=1710–1719|doi=10.1111/j.1095-8649.2007.01445.x}}</ref> 

<gallery mode="packed" heights="150px">
File:Amphiprion ocellaris, puesta.jpg|''A. ocellaris'' male tending to eggs
File:Anemone Fish Eggs.jpg|Clownfish eggs closer to hatching
File:Clownfish growth.webp|Development of ''A. ocellaris'' (above) and ''A. frenatus''
</gallery>

==Captivity==
[[File:Black Clown Fishes (01112491) (6175426265).jpg|thumb|right|Melanistic clownfish in fish tank]]
Clownfish have been popular in the [[Fishkeeping|aquarium trade]] due to their colouration, sociability and longevity. With no predators, they can thrive in captivity without sea anemones.<ref>da Silva, C. R. B.; Hoepner, C. M.; Mercader, M.; Laudet, V.; da Silva, K. B. "The Impact of Popular Film on the Conservation of Iconic Species: Anemonefishes in the Aquarium Trade" in {{harvnb|Laudet|Ravasi|2023}} pp. 223, 227</ref> Between 1997 and 2002, ''A. ocellaris'' was among the most traded species worldwide<ref>{{cite book|author=Taylor, M.|url=http://www.unep.org/pdf/from_ocean_to_aquarium_report.pdf|title=From ocean to aquarium: A global trade in marine ornamental species|author2=Razak, T.|author3=Green, E.|publisher=UNEP world conservation and monitoring centre (WCMC)|year=2003|page=19|access-date=18 April 2013|archive-url=https://web.archive.org/web/20040701215234/http://www.unep.org/pdf/from_ocean_to_aquarium_report.pdf|archive-date=July 1, 2004|name-list-style=amp}}</ref>, while in 2011, the species was the most popularly imported clownfish species in the US, being almost half of individual clownfish, and around three percent of total marine fishes. Other popularly traded clownfishes include ''A. biaculeatus'' and ''A. percula''. Highest amount of individual clownfish were imported from the Philippines and Indonesia.<ref>da Silva, C. R. B.; Hoepner, C. M.; Mercader, M.; Laudet, V.; da Silva, K. B. "The Impact of Popular Film on the Conservation of Iconic Species: Anemonefishes in the Aquarium Trade" in {{harvnb|Laudet|Ravasi|2023}} pp. 225–226</ref><ref>{{cite journal|last1=Rhyne|first1=A. L.|last2=Tlusty|first2=M. F.|last3=Szczebak|first3=J. T.|last4=Holmberg|first4=R. J.|year=2017|title=Expanding our understanding of the trade in marine aquarium animals|journal=PeerJ|volume=5|page=e2949|doi=10.7717/peerj.2949|doi-access=free}}</ref> 

Clownfish are featured in the 2003 [[Disney/Pixar]] film ''[[Finding Nemo]]'' which has been suggested to have led to an increase in the taking and purchase of clownfish.<ref>{{cite web|title=Buying Nemo|website=E The Environmental Magazine|date=29 July 2004|url=https://emagazine.com/buying-nemo/|accessdate=15 May 2025}}</ref><ref>{{cite journal|last1=McClenachan|first1=L|last2=Cooper|first2=A. B.|last3=Carpenter|first3=K. E.|last4=Dulvy|first4=N. K.|year=2012|title=Extinction risk and bottlenecks in the conservation of charismatic marine species|journal=Conservation Letters|volume=5|issue=1|pages=73–80|doi=10.1111/j.1755-263X.2011.00206.x}}</ref> However, a 2017 study found no evidence for an increase in the sales of wild-caught ''A. ocellaris'' or ''A. percula''—the two species in which the title character bears a resemblance to.<ref>{{cite journal|last1=Militz|first1=T. A.|last2=Foale|first2=S|year=2017|title="The "Nemo Effect": Perception and reality of ''Finding Nemo''s impact on marine aquarium fisheries|journal=Fish and Fisheries|volume=18|issue=3|pages=596–606|doi=10.1111/faf.12202}}</ref> Clownfish are also bred in [[aquaculture]]; though it is unknown how their numbers compare to those wild caught specimens. In 2011, ''[[Amphiprion latezonatus|A. latezoneatus]]'', which is [[endemic]] to Australia, was mostly imported to the US from the Philippines, suggesting they were captive bred. Captive clownfish with new colour morphs, known as "designer clownfish", are particularly popular in the trade.<ref>da Silva, C. R. B.; Hoepner, C. M.; Mercader, M.; Laudet, V.; da Silva, K. B. "The Impact of Popular Film on the Conservation of Iconic Species: Anemonefishes in the Aquarium Trade" in {{harvnb|Laudet|Ravasi|2023}} pp. 226, 228</ref>

==Conservation==
As of 2025, 25 of the 28 clownfish species along with the hybrid ''A. leucokranos'' are assessed by the [[IUCN Red List]] as [[Least-concern species|Least-concern]]. Additionally, ''A. latezonatus'' is listed as Data Deficient.<ref>{{cite website|url=https://www.iucnredlist.org/search?query=amphiprion&searchType=species|title=Search for "amphiprion"|accessdate=16 May 2025|website=IUCN Red List of Threatened Species}}</ref> Threats to clownfish populations in the wild include, [[Ocean heat content|ocean warming]] and [[Ocean acidification|acidification]], exploitation for the aquarium trade and human development along the coast;<ref name=Jones287>Jones, G. P.; Srinivasan, M.; Galbraith, G. F.; Berumen, M. L.; Planes, S. "Saving Nemo: Extinction Risk, Conservation Status, and Effective Management Strategies for Anemonefishes" in {{harvnb|Laudet|Ravasi|2023}} pp. 287–288</ref> the first two are linked to [[global warming]].<ref>Schunter, C.; Donelson, J. M.; Munday, P. L.; Ravasi, T. "Resilience and Adaptation to Local and Global Environmental Change" in {{harvnb|Laudet|Ravasi|2023}} pp. 253–254</ref> 
[[File:Pez payaso (Amphiprion bicinctus) en una anémona magnífica (Heteractis magnifica), mar Rojo, Egipto, 2023-04-18, DD 93.jpg|thumb|right|''[[Amphiprion bicinctus|A. bicinctus]]'' which is endemic to the [[Red Sea]]]]
A warmer  ocean could lead to a delay in the development and settling of larvae.<ref>{{cite journal|last1=McLeod|first1=I. M.|last2=Rummer|first2=J. L.|last3=Clark|first3=T. D.|last4=Jones|first4=G. P.|last5=McCormick|first5=M. I.|last6=Wenger|first6=A. S.|last7=Munday|first7=P. L.|year=2013|title=Climate change and the performance of larval coral reef fishes: the interaction between temperature and food availability|journal=Conservation Physiology|volume=1|issue=1|page=cot024|doi=10.1093/conphys/cot024}}</ref> increase their need to forage,<ref>{{cite journal|last1=Nowicki|first1=J. P.|last2=Miller|first2=G. M.|last3=Munday|first3=P. L.|year=2012|title=Interactive effects of elevated temperature and CO2 on foraging behavior of juvenile coral reef fish|journal=Journal of Experimental Marine Biology and Ecology|volume=412|pages=46–51|doi=10.1016/j.jembe.2011.10.020}}</ref> as well as a decreased reproduction.<ref name=Miller2015>{{cite journal|last1=Miller|first1=G. M.|last2=Kroon|first2=F. J.|last3=Metcalfe|first3=S|last4=Mundayi|first4=P. L.|year=2015|title=Temperature is the evil twin: effects of increased temperature and ocean acidification on reproduction in a reef fish|journal=Ecological Applications|volume=25|issue=3|pages=603–620|doi=10.1890/14-0559.1}}</ref> Conversely, a 2015 study suggested that warmer water can increase [[aerobic exercise]] in ''A. biaculeatus'' juveniles.<ref>{{cite journal|last=Donelson|first=J. M.|year=2015|title=Development in a warm future ocean may enhance performance in some species|journal=Journal of Experimental Marine Biology and Ecology|volume=472|pages=119–125|doi=10.1016/j.jembe.2015.07.008}}</ref> Warming also causes the bleaching of their anemone hosts (similar to [[coral bleaching]]); a 2008 study concluded that anemone bleaching has led to lower numbers of clownfish in [[Great Keppel Island]], off [[Queensland]], Australia.<ref>{{cite journal|last1=Jones|first1=A. M.|last2=Gardner|first2=S|last3=Sinclair|first3=W|year=2008|title=Losing ‘Nemo’: bleaching and collection appear to reduce inshore populations of anemonefishes|journal=Journal of Fish Biology|volume=73|issue=3|pages=753–761|doi=10.1111/j.1095-8649.2008.01969.x}}</ref> Another study found that clownfish can actually help their hosts better recover from bleaching.<ref>{{cite journal|last1=Pryor|first1=S. H.|last2=Hill|first2=R|last3=Dixson|first3=D. L.|last4=Fraser|first4=N. J.|last5=Kelaher|first5=B. P.|last6=Scott|first6=A|year=2020|title=Anemonefish facilitate bleaching recovery in a host sea anemone|journal=Scientific Reports|volume=10|page=18586|doi=10.1038/s41598-020-75585-6}}</ref> There is evidence that ocean acidification negatively impacts clownfish larvae's sense of smell and hearing, which consequently reduces their ability to find reefs and anemones and increases predation risk.<ref>{{cite journal|last1=Munday|first1=P. L.|last2=Dixon|first2=D. L.|last3=Donelson|first3=J. M.|last4=Jones|first4=G. P.|last5=Pratchett|first5=M. S.|last6=Devitsina|first6=G. V.|last7=Døving|first7=K. B.|year=2009|title=Ocean acidification impairs olfactory discrimination and homing ability of a marine fish|journal=Proceedings of the National Academy of Science|volume=106|issue=6|pages=1848–1852|doi=10.1073/pnas.0809996106}}</ref><ref>{{cite journal|last1=Simpsons|first1=S. D.|last2=Munday|first2=P. L.|last3=Wittenrich|first3=M. L.|last4=Manassa|first4=R|last5=Dixson|first5=D. L.|last6=Gagliano|first6=M|last7=Yan|first7=H. Y.|year=2011|title=Ocean acidification erodes crucial auditory behaviour in a marine fish|journal=Biology Letters|volume=7|issue=6|page=917–920|doi=10.1098/rsbl.2011.0293}}</ref><ref name=Jones287/> Conversely, a 2013 study found that higher levels of [[carbon dioxide]] in the waters increased [[fecundity]] in adults, but also led to smaller yolks for larvae.<ref>{{cite journal|last1=Miller|first1=G. M.|last2=Watson|first2=S-A|last3=McCormick|first3=M. I.|last4=Munday|first4=P. L.|year=2013|title=Increased CO2 stimulates reproduction in a coral reef fish|journal=Global Change Biology|volume=19|issue=10|pages=3037–3045|doi=10.1111/gcb.12259}}</ref> In addition, acidification can negatively affect the quality of larvae when combined with warmer temperatures.<ref name=Miller2015/>

Wild clownfish are commonly collected to supply the aquarium trade. Factors that contribute to this include their popularity, market value, and ease of capture, particularly due to their shallow water habitat.<ref name=Jones287/> A 2005 study of an area in the Philippines found that clownfish and anemones were suspect to overfishing, being 60 percent of total catch; clownfish in particular decreased in population density by 80 percent.<ref>{{cite journal|last1=Shuman|first1=C. S.|last2=Hodgson|first2=G|last3=Ambrose|first3=R. F.|year=2005|title=Population impacts of collecting sea anemones and anemonefish for the marine aquarium trade in the Philippines|journal=Coral Reefs|volume=24|pages=564–573|doi=10.1007/s00338-005-0027-z}}</ref> A 2014 study also concluded that exploitation has led to a decrease in ''O. ocellaris'' and associated anemones around the [[Spermonde Archipelago]], Indonesia.<ref>{{cite journal|last1=Madduppa|first1=H. H.|last2=von Juterzenka|first2=K|last3=Syakir|first3=M|last4=Kochzius|first4=M|year=2014|title=Socio-economy of marine ornamental fishery and its impact on the population structure of the clown anemonefish Amphiprion ocellaris and its host anemones in Spermonde Archipelago, Indonesia|journal=Ocean & Coastal Management|volume=100|issue=2|pages=41–50|doi=10.1016/j.ocecoaman.2014.07.013}}</ref> Frisch and colleagues (2019) found that during a pause on collecting in [[Keppel Bay Islands National Park|Keppel Islands]], Australia, the recovery rate for clownfish species was relatively slow over a ten year period.<ref>{{cite journal|last1=Frisch|first1=A. J.|last2=Hobbs|first2=J-P. A.|last3=Hansen|first3=S. T.|last4=Williamson|first4=D. H.|last5=Bonin|first5=M. C.|last6=Jones|first6=G. P.|last7=Rizzari|first7=J. R.|year=2019|title=Recovery potential of mutualistic anemone and anemonefish populations|journal=Fisheries Research|volume=218|pages=1–9|doi=10.1016/j.fishres.2019.04.018}}</ref> Urban development along the coast can effect clownfish habitat via increased sedimentation, algae growth and [[dredging]].<ref name=Jones287/> Sedimentation in particular appears have negative effects on larval development,<ref>{{cite journal|last1=Wenger|first1=A. S.|last2=McCormick|first2=M. I.|last3=Endo|first3=G. G. K.|last4=McLeod|first4=I. M.|last5=Kroon|first5=F|last6=Jones|first6=G. P.|year=2014|title=Suspended sediment alters larval development in a coral reef fish|journal=Journal of Experimental Biology|volume=217|issue=7|pages=1122–1128|doi=10.1242/jeb.094409}}</ref> gill function<ref>{{cite journal|last1=Hess|first1=S|last2=Wenger|first2=A. S.|last3=Ainsworth|first3=T. D.|last4=Rummer|first4=J. L.|year=2015|title=Exposure of clownfish larvae to suspended sediment levels found on the Great Barrier Reef: Impacts on gill structure and microbiome|journal=Scientific Reports|volume=5|page=10561|doi=10.1038/srep10561}}</ref> and anti-predator behaviour.<ref>{{cite journal|last1=Hess|first1=S|last2=Allan|first2=B. J. M.|last3=Hoey|first3=A. S.|last4=Jarrold|first4=M. D.|last5=Wenger|first5=A. S.|last6=Runner|first6=J. L.|year=2019|title=Enhanced fast-start performance and anti-predator behaviour in a coral reef fish in response to suspended sediment exposure alters larval development in a coral reef fish|journal=Coral Reefs|volume=38|issue=1|pages=103–108|doi=10.1007/s00338-018-01757-6}}</ref> Declines in the numbers of certain clownfish species and their anemones hosts have been linked to coastal development in certain areas.<ref>{{cite journal|last1=Hayashi|first1=K|last2=Tachihara|first2=K|last3=Reimer|first3=J. D.|year=2019|title= Loss of natural coastline influences species diversity of anemonefish and host anemones in the Ryuku Archipelago|journal=Aquatic Conservation: Marine and Freshwater Ecosystems|volume=31|issue=1|pages=15–27|doi=10.1002/aqc.3435}}</ref><ref>{{cite journal|last1=Howell|first1=J|last2=Goulet|first2=T. L.|last3=Goulet|first3=D|year=2016|title= Anemonefish musical chairs and the plight of the two-band anemonefish, ''Amphiprion bicinctus''|journal=Environmental Biology of Fishes|volume=99|pages=873–886|doi=10.1007/s10641-016-0530-9}}</ref>

The establishment of [[marine protected area]]s has benefited clownfish due to prohibition on collecting, as they appear to be more numerous in these areas.<ref name=Jones291>Jones, G. P.; Srinivasan, M.; Galbraith, G. F.; Berumen, M. L.; Planes, S. "Saving Nemo: Extinction Risk, Conservation Status, and Effective Management Strategies for Anemonefishes" in {{harvnb|Laudet|Ravasi|2023}} pp. 291–293</ref> There are particularly important for protecting biodiversity hotspots where multiple clownfish species live, such as [[Solitary Islands Marine Park]], Australia.<ref name=Jones291/><ref>{{cite journal|last1=Scott|first1=A|last2=Malcolm|first2=H. A.|last3=Damiano|first3=C|last4=Richardson|first4=D. L.|year=2011|title=Long-term increases in abundance of anemonefish and their host sea anemones in an Australian marine protected area|journal=Marine and Freshwater Research|volume=62|issue=2|pages=187–196|doi=10.1071/MF10323}}</ref> Marine parks are important for protecting endemic species, but do not appear to increase the recovery of species that have had wide-ranging declines.<ref name=Jones291/>

==Notes==
{{Notelist}}

==References==
{{reflist}}

===Bibliography===
*{{cite book|last1=Allen|first1=G. R.|year=1975b|title=Anemonefishes: Their Classification and Biology|publisher=T. F. H. Publications Ltd|edition=2nd|isbn=0-87666-001-4}}
*{{cite book|last1=Fautin|first1=D. G.|last2=Allen|first2=G. R.|year=1992|title=Field Guide to Anemonefishes and Their Host Sea Anemones|publisher=Western Australian Museum|isbn=0-7309-5216-9}}
*{{cite book|editor1-last=Frédérich|editor1-first=B|editor2-last=Parmentier|editor2-first=E|year=2016|title=Biology of Damselfishes|publisher=CRC Press|isbn=9781482212099 |doi=10.1201/9781315373874}}
*{{cite book|editor1-last=Laudet|editor1-first=V|editor2-last=Ravasi|editor2-first=T|year=2023|title=Evolution, Development and Ecology of Anemonefishes|publisher=CRC Press|isbn=978-0367645816|doi=10.1201/9781003125365}}

==External links==
{{Commons category|Amphiprioninae}}
{{Wikispecies|Amphiprioninae}}
* {{in lang|de}} [http://www.torsten-ernst.de/modules.php?name=Gallery&file=categories&cat_id=13 Photo Gallery of ''Amphiprion ocellaris'' and their eggs] {{Webarchive|url=https://web.archive.org/web/20210313145247/http://www.torsten-ernst.de/modules.php?name=Gallery&file=categories&cat_id=13 |date=13 March 2021 }}
* [https://web.archive.org/web/20080724075222/http://www.montereybayaquarium.org/efc/efc_splash/splash_animals_clownfish.aspx Monterey Bay Aquarium: Video and information]
* [https://web.archive.org/web/20100609112916/http://www.tommyschultz.com/component/searchimage/clown-fish-best/1.html Clown Fish underwater photography gallery]

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[[Category:Pomacentridae]]
[[Category:Symbiosis]]
[[Category:Articles containing video clips]]
[[Category:Ray-finned fish subfamilies]]