Amphiprioninae

A pair of pink anemonesh (Amphiprion perideraion) in their

anemone home.

Ocellaris clownsh nestled in a magnicent sea anemone

(Heteractis magnica)

Clownsh or anemonesh are shes from the subfamily

Amphiprioninae in the family Pomacentridae. Thirty
species are recognized, one in the genus Premnas, while
the remaining are in the genus Amphiprion. In the wild,
they all form symbiotic mutualisms with sea anemones.
Depending on species, Clownsh are overall yellow, orange, or a reddish or blackish color, and many show white
bars or patches. The largest can reach a length of 18 centimetres (7.1 in), while the smallest barely can reach 10
centimetres (3.9 in).
Clown sh swimming movements

2 Diet
1

Ecology and habitat

Clownsh are omnivorous and can feed on undigested

food from their host anemones, and the fecal matter
from the clownsh provides nutrients to the sea anemone.
Clownsh primarily feed on small zooplankton from the
water column, such as copepods and tunicate larvae, with
a small portion of their diet coming from algae, with
the exception of Amphiprion perideraion, which primarily feeds on algae.[1][2] They may also consume the tenta-

Clownsh are native to warmer waters of the Indian and

Pacic oceans, including the Great Barrier Reef and the
Red Sea. While most species have restricted distributions, others are widespread. Clownsh live at the bottom
of shallow seas in sheltered reefs or in shallow lagoons.
There are no clownsh in the Atlantic.
1

5 PARENTAL INVESTMENT
haviours and movements such as wedging and switching. Aeration of the host anemone tentacles allows for
benets to the metabolism of both partners, mainly by
increasing anemone body size and both clownsh and
anemone respiration.[9]
Clownsh, Cardinalsh and certain damselsh are among
the few species of sh that can avoid the potent poison of
a sea anemone. There are several theories about how they
can survive the sea anemone poison:

A clownsh swimming.

The mucus coating of the sh may be based on

sugars rather than proteins. This would mean that
anemones fail to recognize the sh as a potential
food source and do not re their nematocysts, or
sting organelles.
The coevolution of certain species of clownsh with
specic anemone host species and may have acquired an immunity to the nematocysts and toxins
of their host anemone. Experimentation has shown
that Amphiprion percula may develop resistance to
the toxin from Heteractis magnica, but it is not totally protected, since it was shown experimentally to
die when its skin, devoid of mucus, was exposed to
the nematocysts of its host.[10]

4 Reproduction
Video of a clownsh swimming around an anemone.

In a group of clownsh, there is a strict dominance hierarchy. The largest and most aggressive female is found
cles of their host anemone.[3]
at the top. Only two clownsh, a male and a female, in a
group reproduce through external fertilization. Clownsh
are sequential hermaphrodites, meaning that they develop
into males rst, and when they mature, they become fe3 Symbiosis and mutualism
males. If the female clownsh is removed from the group,
such as by death, one of the largest and most dominant
Clownsh and sea anemones have a symbiotic, mutu- males will become a female. The remaining males will
alistic relationship, each providing a number of bene- move up a rank in the hierarchy.
ts to the other. The individual species are generally
Clownsh lay eggs on any at surface close to their host
highly host specic, and especially the genera Heteractis
anemones. In the wild, clownsh spawn around the time
and Stichodactyla, and the species Entacmaea quadriof the full moon. Depending on the species, clownsh
color are frequent clownsh partners. The sea anemone
can lay hundreds or thousands of eggs. The male parent
protects the clownsh from predators, as well as proguards the eggs until they hatch about six to ten days later,
viding food through the scraps left from the anemones
typically two hours after dusk.
meals and occasional dead anemone tentacles. In return, the clownsh defends the anemone from its predators, and parasites.[4][5] The anemone also picks up nutrients from the clownshs excrement, and functions as 5 Parental investment
a safe nest site.[6] The nitrogen excreted from clownsh increases the amount of algae incorporated into the Most clownsh are protandrous hermaphrodites, meantissue of their hosts, which aids the anemone in tissue ing they alternate between the male and female sexes at
growth and regeneration.[2] It has been theorized that the some point in their lives. Anemonesh colonies usually
clownsh use their bright coloring to lure small sh to the consist of the reproductive male and female and a few
anemone,[7] and that the activity of the clownsh results in juveniles, who help tend the colony.[11] Although multigreater water circulation around the sea anemone.[8] Stud- ple males co-habit an environment with a single female,
ies on anemonesh have found that clownsh alter the polygamy does not occur and only the adult pair exhibit
ow of water around sea anemone tentacles by certain be- reproductive behavior. However, if the largest female

3
dies, the social hierarchy shifts with the breeding male
exhibiting protandrous sex reversal to become the breeding female. The largest juvenile will then become the
new breeding male after a period of rapid growth.[12]
The existence of protandry in clownsh may rest on the
case that non-breeders modulate their phenotype in a way
that causes breeders to tolerate them. This strategy prevents conict by reducing competition between the males
for one female. For example, by purposefully modifying their growth rate to remain small and submissive, the
juveniles in a colony present no threat to the tness of
the adult male, thereby protecting themselves from being
evicted by the dominant sh.[13]
The reproductive cycle of clownsh is often correlated
with the lunar cycle. Rates of spawning for clownsh
peak at approximately the rst and third quarters of the
moon. The timing of this spawn means that the eggs will
hatch around the full moon or new moon periods. One
explanation for this lunar clock is that spring tides produce the highest tides during full or new moons. Nocturnal hatching during high tide may reduce predation by
allowing for a greater capacity for escape. Namely, the
stronger currents and greater water volume during high
tide protects the hatchlings by eectively sweeping them
to safety. Before spawning, clownsh exhibit increased
rates of anemone and substrate biting, which help prepare
and clean the nest for the spawn.[12]

sh bred in captivity, while the majority are captured

from the wild,[15][16] accounting for decreased densities
in exploited areas.[17] Public aquaria and captive breeding programs are essential to sustain their trade as marine ornamentals, and has recently become economically
feasible.[18][19] It is one of a handful of marine ornamentals whose complete life cycle has been closed in captivity. Members of some clownsh species, such as the maroon clownsh, become aggressive in captivity; others,
like the false percula clownsh, can be kept successfully
with other individuals of the same species.
When a sea anemone is not available in an aquarium,
the clownsh may settle in some varieties of soft corals,
or large polyp stony corals. Once an anemone or coral
has been adopted, the clownsh will defend it. As there
is less pressure to forage for food in an aquarium, it is
common for clownsh to remain within 2-4 inches of
their host for their entire lifetime. Clownsh, however,
are not obligately tied to hosts, and can survive alone in
captivity.[20][21]

7 Taxonomy
Genus Amphiprion:[22]
Amphiprion akallopisos Skunk clownsh

In terms of parental care, male clownsh are often the

caretakers of eggs. Before making the clutch, the parents often clear an oval sized clutch varying in diameter
for the spawn. Fecundity, or reproductive rate, of the females usually ranges from 600 to 1500 eggs depending on
the size of the female. In contrast to most animal species,
the female only occasionally takes responsibility for the
eggs, with males expending most of the time and eort.
Male clownsh care for their eggs by fanning and guarding them for 6 to 10 days until they hatch. Studies have
shown that, in general, eggs develop more rapidly in a
clutch when males fanned properly and that fanning represents a crucial mechanism of successfully developing
eggs. This suggests that males have the ability to control
the success of hatching an egg clutch by investing dierent amounts of time and energy towards the eggs. For example, a male could choose to fan less in times of scarcity
or fan more in times of abundance. Furthermore, males
display increased alertness when guarding more valuable
broods, or eggs in which paternity was guaranteed. Females, on the other hand, display generally less preference for parental behavior than males. All these suggest
that males have increased parental investment towards the
eggs compared to females.[14]

Amphiprion akindynos
anemonesh

Marine ornamentals

Clownsh make up 43% of the global marine ornamental trade, and 25% of the global trade comes from

Barrier

reef

Amphiprion allardi Twobar anemonesh

Amphiprion barberi
Amphiprion bicinctus Twoband anemonesh
Amphiprion chagosensis Chagos anemonesh
Amphiprion
anemonesh

chrysogaster

Mauritian

Amphiprion
anemonesh

chrysopterus

Orange-n

Amphiprion clarkii Yellowtail clownsh

Amphiprion ephippium Saddle anemonesh
Amphiprion frenatus Tomato clownsh
Amphiprion fuscocaudatus
anemonesh

Seychelles

Amphiprion latezonatus
Anemonesh

Wide-band

Amphiprion latifasciatus
anemonesh

Madagascar

Amphiprion leucokranos
anemonesh

Whitebonnet

Amphiprion
anemonesh

Whitesnout

mccullochi

Amphiprion melanopus Fire clownsh

Amphiprion nigripes Maldive anemonesh

10
Amphiprion ocellaris Clown anemonesh
Amphiprion omanensis Oman anemonesh
Amphiprion pacicus Pacic anemonesh
Amphiprion percula Orange clownsh
Amphiprion perideraion Pink skunk clownsh
Amphiprion polymnus Saddleback clownsh
Amphiprion rubrocinctus Red Anemonesh
Amphiprion sandaracinos Yellow clownsh
Amphiprion sebae Sebae anemonesh
Amphiprion thiellei Thielles anemonesh
Amphiprion tricinctus Three-band anemonesh
Genus Premnas:[23]
Premnas biaculeatus Maroon clownsh

In popular culture

In the 2003 animated lm Finding Nemo, the main character is a clownsh. The species depicted in the lm is A.
ocellaris.[4]

REFERENCES

Maroon clownsh (Premnas biaculeatus), here with

a tang, is the only member of the genus Premnas,
and the largest species of clownsh.
Clownsh are initially male; the most dominant sh
in a group becomes female.

10 References
[1] Fautin, Daphne; Gerald Allen (1997). Field Guide to
Anemone Fishes and Their Host Sea Anemones (2 ed.).
Perth, Australia: Western Australian Museum. ISBN
978-0-7309-8365-1.
[2] Porat, D.; Chadwick-Furman, N.E. (2005). Eects of
anemonesh on giant sea anemones: Ammonium uptake,
zooxanthella content and tissue regeneration. Marine
and Freshwater Behaviour and Physiology 29 (1): 4351.
doi:10.1080/10236240500057929.
[3] Fautin, D.G.; Guo, C.; Hwang, J.S. (1995). Costs and
benets of the symbiosis between the anemoneshrimp
Periclimenes brevicarpalis and its host Entacmaea quadricolor.. Marine Ecology Progress Series 129: 7784.
doi:10.3354/meps129077.
[4] Clown Anemonesh. Nat Geo Wild : Animals. National
Geographic Society. Retrieved 2011-12-19.
[5] Amphiprioninae at the Encyclopedia of Life

Gallery
Allards clownsh (Amphiprion allardi).
Maldive anemonesh (Amphiprion nigripes). It is
sometimes called the Black-nned anemonesh due
to its black pelvic and anal ns.
Pink skunk clownsh (Amphiprion perideraion).
Saddleback clownsh (Amphiprion polymnus) o
Sulawesi, Indonesia.
Yellow clownsh (Amphiprion sandaracinos) and
sea anemone o Sulawesi, Indonesia.
Yellowtail clownsh (Amphiprion clarkii) with sea
anemone.
Orange-n anemonesh (Amphiprion chrysopterus)
is one of the few anemonesh with a white tail.
Twoband anemonesh (Amphiprion bicinctus). It is,
as suggested by its alternative common name Red
Sea clownsh, almost entirely restricted to the Red
Sea.
Clown anemonesh (Amphiprion ocellaris) occurs in
a 'normal' orange and a melanistic blackish variant.
The exact taxonomic status of the latter is unclear.
Fire clownsh (Amphiprion melanopus) and sea
anemone at the Great Barrier Reef.

[6] Holbrook, S. J. and Schmitt,R. J. Growth, reproduction

and survival of a tropical sea anemone (Actiniaria): benets of hosting anemonesh, 2005, cited in
[7] Clown Anemoneshes,
Amphiprion ocellaris".
Marinebio.
The MarineBio Conservation Society.
Retrieved 2011-12-19.
[8] Szczebak, Joseph T.; Raymond P. Henry; Fuad A.
Al-Horani; Nanette E. Chadwick (2013). Anemonesh
oxygenate their anemone hosts at night.
Journal of Experimental Biology 216 (9): 970976.
doi:10.1242/jeb.075648.
[9] Joseph T. Szczebak, Raymond P. Henry, Fuad A.
Al-Horani, Nanette E. Chadwick (2012-11-03).
Anemonesh oxygenate their anemone hosts at night.
The Journal of Experimental Biology.
Retrieved
2013-09-15.
[10] Mebs, D. 1994. Anemonesh symbiosis: Vulnerability
and Resistance of Fish to the Toxin of the Sea Anemone.
Toxicon. Vol. 32(9):10591068.
[11] Stephanie Boyer. Clown Anemosh. Florida Museum
of Natural History. Retrieved 2013-09-15.
[12] Robert M. Ross (1978-02-10). Reproductive Behavior
of the Anemonesh Amphiprion melanopus on Guam.
Copeia. Retrieved 2013-09-15.
[13] Peter Buston (2004-08-18). Does the presence of nonbreeders enhance the tness of breeders? An experimental analysis in the clown anemonesh Amphiprion percula. Springer-Verlag. Retrieved 2013-09-15.

[14] Swagat Ghosh, T. T. Ajith Kumar, T. Balasubramanian

(2011-08-04). Determining the level of parental care relating fanning behavior of ve species of clownshes in
captivity. Indian Journal of Geo-Marine Sciences. Retrieved 2013-09-15.
[15] Dhaneesh, K.V.; R. Vinoth, Swagat Gosh, M. Gopi, T.T.
Ajith Kumar, T. Balasubramanian (2013). Sundaresan, J.,
ed. Hatchery Production of Marine Ornamental Fishes:
An Alternate Livelihood Option for the Island Community at Lakshadweep. Climate Change and Island and
Coastal Vulnerability (Capital Publishing Company) 17:
253265. doi:10.1007/978-94-007-6016-5_17.
[16] Taylor, M., Green, E. and Razak, T. (2003). From
ocean to aquarium: A global trade in marine ornamental
species. UNEP world conservation and monitoring centre
(WCMC). pp. 164. Retrieved 18 April 2013.
[17] Shuman, Craig; Gregor Hodgson; Richard F. Ambrose (2005). Population impacts of collecting sea
anemones and anemonesh for the marine aquarium
trade in the Philippines. Coral Reefs 24: 564573.
doi:10.1007/s00338-005-0027-z.
[18] Watson, Craig; Jeery Hill (2006). Design criteria for recirculating, marine ornamental production systems. Aquacultural Engineering 34 (3): 157162.
doi:10.1016/j.aquaeng.2005.07.002.
[19] Hall, Heather; Douglas Warmolts (2003). 23. In
James C. Cato, Christopher L. Brown. Marine Ornamental Species: Collection, Culture and Conservation. WileyBlackwell. pp. 303326. ISBN 978-0-8138-2987-6.
[20] Daphne Gail Fautin (1991). The anemonesh symbiosis:
what is known and what is not (PDF). Symbiosis 10: 23
46.
[21] Ronald L. Shimek (2004). Marine Invertebrates. Neptune
City, NJ: T.F.H. Publications. p. 83. ISBN 978-1890087-66-1.
[22] Froese, Rainer, and Daniel Pauly, eds. (2011). Species of
Amphiprion in FishBase. December 2011 version.
[23] Froese, Rainer, and Daniel Pauly, eds. (2011). Species of
Premnas in FishBase. December 2011 version.

11

External links

Clownsh species proles on AquariumDomain.

(German) Photo Gallery of Amphiprion ocellaris
and their eggs
Monterey Bay Aquarium: Video and information
Clown Fish underwater photography gallery
Aquaticcommunity.com
Tolweb.org

12

12
12.1

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

Text and image sources, contributors, and licenses

Text

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12.2

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File:Amphiprion_ocellaris_(Clown_anemonefish)_in_Heteractis_magnifica_(Sea_anemone).jpg Source: http://upload.wikimedia.

org/wikipedia/commons/e/e5/Amphiprion_ocellaris_%28Clown_anemonefish%29_in_Heteractis_magnifica_%28Sea_anemone%29.
jpg License: CC BY-SA 3.0 Contributors: Own work Original artist: Nhobgood Nick Hobgood
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talk:Nhobgood'>talk</a>) Nick Hobgood

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