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Clownfish

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Clownfish
Amphiprion ocellaris
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Blenniiformes
Family: Pomacentridae
Subfamily: Amphiprioninae
Allen, 1975
Genus: Amphiprion
Bloch & Schneider, 1801
Synonyms

Clownfishes or anemonefishes are saltwater fishes 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 reefs. They have a distinctive colouration typically consisting of white vertical bars on a red, orange, yellow, brown or black background. Clownfish have developed a mutualistic symbiotic relationship with sea anemones, 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 planktonic 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 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 fertilises 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 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 areas, where collecting is forbidden. Other threats to populations include global warming which causes ocean warming and acidification.

Taxonomy

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Clownfish are damselfish (family Pomacentridae) in the genus Amphiprion (Greek amphi, 'on both sides' and prion 'saw'), which was coined by Marcus Elieser Bloch and Johann Gottlob Theaenus Schneider in 1801.[1][2] Georges Cuvier considered the maroon clownfish (specific name biaculeatus) morphologically different enough to be placed in its own genus Premnas in 1817.[3] 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.[1][4] In 1975, ichthyologist Gerald R. Allen placed clownfishes in their own subfamily Amphiprioninae.[5] A 2009 study suggested creating the tribe Amphiprionini for clownfishes and moving them to the subfamily Pomacentrinae.[6]

Phylogeny

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The clownfish lineage diverged from other living damselfishes around 35 million years ago (mya) during the late Eocene,[4] and the most recent common ancestor is dated around 10.5 mya during the Early Miocene[7] in the waters of the Malay Archipelago.[8] Clownfishes experienced an increase in species diversification starting around 5 mya,[4] with two major adaptive radiations; one centred on the Malay Archipelago and later in the waters of the western Indian Ocean.[8] High amounts of interbreeding between species occurred and the most recently diverged species are descended from hybrids.[9] Clownfish speciation has been linked to their sea anemone hosts, species of which can be found in different habitats and thus drove ecological separation.[10][7]

There are 28 living species of clownfish and two hybrids forms, Amphiprion leucokranos (white-bonnet anemonefish) and Amphiprion thiellei (Thielle’s anemonefish).[11] In 1972, Allen listed five major clades or complexes based on morphology; percula, akallopisos, ephippium, polymnus and clarkii with A. biaculeatus being in a monotypic clade close to percula.[12]. 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.[9] A 2021 study placed A. chrysopterus and members of the clade polymnus in the Indian clade,[4] while a 2025 study found the percula clade to be paraphyletic.[7]

The following cladogram of the 28 clownfish species is based on a genetic study by Gaboriau and colleagues (2025):[7][a]

Amphiprion

A. biaculeatus (Maroon clownfish or Spinecheek anemonefish)

A. percula (Orange clownfish)

A. ocellaris (False clownfish)

A. latezoneatus (Wide-band anemonefish)

A. perideraion (Pink skunk anemonefish)

A. pacificus (Pacific anemonefish)

A. akallopisos (Skunk anemonefish)

A. sandaracinos (Yellow skunk anemonefish)

A. tricinctus (Three-band anemonefish)

A. clarkii (Yellow-tail anemonefish)

A. barberi (Barber’s anemonefish)

A. frenatus (Tomato anemonefish)

A. ephippium (Saddle anemonefish)

A. melanopus (Fire anemonefish)

A. rubrocinctus (Red anemonefish)

A. chrysopterus (Orange-fin anemonefish)

A. mccullochi (White-snout anemonefish)

A. akindynos (Barrier reef anemonefish)

A. sebae (Sebae anemonefish)

A. polymnus (Saddleback anemonefish)

A. nigripes (Maldive anemonefish)

A. chagosensis (Chagos anemonefish)

A. chrysogaster (Mauritian anemonefish)

A. fuscocaudatus (Seychelles anemonefish)

A. bicinctus (Two-band anemonefish)

A. omanensis (Oman anemonefish)

A. allardi (Two-bar anemonefish)

A. latifasciatus (Madagascar anemonefish)

Characteristics

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Clownfish vary in size; A. biaculeatus can reach 160 mm (6.3 in) long while A. percula reaches only 80 mm (3.1 in).[13]. Females are larger than males and the smallest individuals in a group are only 6 mm (0.24 in).[14][15] 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 oral and pharyngeal jaws but absent on the palate, and may be conical or chisel-shaped.[16] They have saw-shaped edges along the operculum (gill covering) and suborbital area, which is the source of their genus name. The dorsal fin has 10 spines and 14–20 soft rays;[17] A. ocellaris has a large recess between the spines and soft rays, while in A. ephippium they are mostly continuous.[18] Rays number 15–21 in the pectoral fins, five in the pelvic fins, 11–15 in the anal fin, and 14–15 in the caudal fin.[19]

Colour patterns

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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 pigment cells, black and brown by melanophores and the white bars by iridophores. 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 A. melanopus, A. polymnus and A. clarkii.[18][20] Numerous 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.[21]

Merilata and colleagues (2018) found that clownfish with only one or no vertical bars tend to be more 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 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.[22] Salis and colleagues (2018) suggested that the bars function in species recognition, finding little overlap between species with the same bar numbers within various clownfish communities.[18] A 2024 study found evidence for this function as A. ocellaris can distinguish between individuals of different bar numbers.[23] Merilata and colleagues (2018) dispute this, noting the geographic and ecological overlap between the similar-looking A. percula and A. ocellaris.[22]

Distribution and habitat

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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.[24][25] Some species are more widespread than others, and some live only around islands or archipelagos. 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.[24] Clownfishes are limited by the distribution of their sea anemone hosts. Hence they are found mainly at the photic zones where there is more photosynthetic zooxanthellae for their hosts to depend on. This includes coral reefs and surrounding areas.[26] 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).[27]

Behaviour and ecology

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Video of a A. melanopus swimming around an anemone

Feeding

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Clownfish are omnivorous, and mostly feed on planktonic food such as copepods and larval tunicates. 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.[28][29] 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 20 m2 (220 sq ft) of ocean floor away from their anemone. Otherwise they are restricted to feeding in the water column above their host.[30] The dominant pair in a clownfish group feed further from the anemone than the smaller subordinates.[29]

Mutualism

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Further information: Symbiosis and Mutualism (biology)

Clownfish have a mutualistic symbiotic relationship with sea anemones.[14][31] They acclimate themselves to their hosts by touching, nipping and fanning the tentacles over a period of minutes to days.[32] 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 oral disc.[26][14] 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.[14] A less important benefit for clownfish is nourishment from the discharged waste and parasites.[26][14]

A. clarkii with an 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.[26] Nevertheless, clownfish contribute to the survival of their hosts by guarding from anemone-eating fish such as the butterflyfish species Chaetodon lunula.[26][33] 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.[31][34] A 2005 study found that anemone grew and regenerated faster with clownfish them without, and attributed this to ammonium from clownfish waste.[35]

A total of ten sea anemone species are used by clownfish as hosts: Radianthus malu, R. crispa, R. magnifica, Stichodactyla mertensii, S. haddoni, S. gigantea', Cryptodendrum adhaesivum, Entacmaea quadricolor, Heteractis aurora and Macrodactyla doreensis. Some clownfish are generalist in their choice of hosts while others are more specialised. A. clarkii is the most generalised species and utilises all ten anemone species, while nine — A. frenatus, A. chagosensis, A. pacificus, A. fuscocaudatus, A. latifasciatus, A. mccullochi, A. nigripes, 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.[36]

Their ability to avoid being stung is attributed to their mucus coating.[26] There is evidence that clownfish mucus mimics the molecules or bacteria of anemone mucus and lacks trigger for the anemone’s nematocysts (stinging barbs). Mucus thickness may also play a role, but this is not clear.[32] 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.[26] 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".[37] Roux and colleagues (2019) found evidence that clownfish exchange microbiota with their anemone hosts.[38]

Social structure

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Group of 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.[39] 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.[39][40] In A. percula, the number of non-breeders ranges from zero to four, with group size depending on anemone size,[41] as well as the size of the female as larger females allow for more members without unbalancing the size ratio between them.[42] Members of a group are unrelated.[43]

As protandrous sequential hermaphrodites, 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.[39][40] New fish that join the group rank at the bottom.[40] 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.[44] The dominant pair controls membership of the group and will drive away individuals when the anemone gets too full,[41] particularly those that are close to them in size. Thus newcomers must remain smaller than their immediate superior to avoid getting evicted.[40] 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.[45] Clownfish appear to produce sounds via the jaws, which is amplified by the swim bladder.[46][47]

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.[48] 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.[23] Conversely, A clarkii was observed to attack individuals of other species more than those of its own.[49]

  • A. clarkii dominant chasing subordinate while producing aggressive sounds
  • A. frenatus subordinate head-shaking while producing submissive sounds

Reproduction and lifecycle

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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 fertilise. Clownfish lay up to a thousand eggs, which are conical in shape, 3–4 mm (0.12–0.16 in) long and stick to the rocky substrate by bundles of short fibres.[50][51] The male tends to the fertilised eggs, cleaning and guarding them as well as fanning them with his pectoral fins.[52]

Incubation lasts six to seven days.[52] 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.[51] 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.[53][54] Larval clownfish are initially transparent, expect for the eyes, yolk-sac and some pigment spots.[53] Over time they begin to metamorphosise; growing in size and developing their fins, sensory and internal organs, notochord flexion and colouration.[54] Clownfish larvae can disperse widely across open ocean; A. omanesis has been recorded travelling over 400 km (250 mi) along ocean currents.[55]

As they enter the juvenile stage, clownfish begin settle to the ocean floor and find an anemone host,[54] while transitioning to a more diurnal lifestyle.[56] Juveniles continue to grow and develop their adult colouration,[54][18] but cannot produce gametes until they ascend to dominance within a group.[54] The dominant male's gonads produce sperm but also possess dormant ovarian cells. When transitioning into a female, the gonads switch to producing ovaries.[57] 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.[58] Clownfish can live for over 20 years.[59] 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.[60]

  • A. ocellaris male tending to eggs
    A. ocellaris male tending to eggs
  • Clownfish eggs closer to hatching
    Clownfish eggs closer to hatching
  • Development of A. ocellaris (above) and A. frenatus
    Development of A. ocellaris (above) and A. frenatus

Captivity

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Melanistic clownfish in fish tank

Clownfish have been popular in the aquarium trade due to their colouration, sociability and longevity. With no predators, they can thrive in captivity without sea anemones.[61] Between 1997 and 2002, A. ocellaris was among the most traded species worldwide[62], 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.[63][64]

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.[65][66] 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.[67] Clownfish are also bred in aquaculture; though it is unknown how their numbers compare to those wild caught specimens. In 2011, 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.[68]

Conservation

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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. Additionally, A. latezonatus is listed as Data Deficient.[69] Threats to clownfish populations in the wild include, ocean warming and acidification, exploitation for the aquarium trade and human development along the coast;[70] the first two are linked to global warming.[71]

A. bicinctus which is endemic to the Red Sea

A warmer ocean could lead to a delay in the development and settling of larvae.[72] increase their need to forage,[73] as well as a decreased reproduction.[74] Conversely, a 2015 study suggested that warmer water can increase aerobic exercise in A. biaculeatus juveniles.[75] 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.[76] Another study found that clownfish can actually help their hosts better recover from bleaching.[77] 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.[78][79][70] 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.[80] In addition, acidification can negatively affect the quality of larvae when combined with warmer temperatures.[74]

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.[70] 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.[81] A 2014 study also concluded that exploitation has led to a decrease in O. ocellaris and associated anemones around the Spermonde Archipelago, Indonesia.[82] Frisch and colleagues (2019) found that during a pause on collecting in Keppel Islands, Australia, the recovery rate for clownfish species was relatively slow over a ten year period.[83] Urban development along the coast can effect clownfish habitat via increased sedimentation, algae growth and dredging.[70] Sedimentation in particular appears have negative effects on larval development,[84] gill function[85] and anti-predator behaviour.[86] Declines in the numbers of certain clownfish species and their anemones hosts have been linked to coastal development in certain areas.[87][88]

The establishment of marine protected areas has benefited clownfish due to prohibition on collecting, as they appear to be more numerous in these areas.[89] There are particularly important for protecting biodiversity hotspots where multiple clownfish species live, such as Solitary Islands Marine Park, Australia.[89][90] Marine parks are important for protecting endemic species, but do not appear to increase the recovery of species that have had wide-ranging declines.[89]

Notes

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  1. ^ Common names are taken from[11]

References

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  2. ^ Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in Frédérich & Parmentier 2016 pp. 303, 306
  3. ^ Colleye, O.; Iwata, E.; Parmentier, E. "Clownfishes" in Frédérich & Parmentier 2016 p. 306
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Bibliography

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