Contents 1 Distribution and habitats 2 Diet 3 Symbiosis and mutualism 4 Reproduction 5 Parental investment 6 In the aquarium 7 Taxonomy 8 In popular culture 9 Gallery 9.1 Clownfish 9.2 A. clarkii complex 9.3 Tomato complex 9.4 Saddleback complex 9.5 Skunk complex 9.6 Premnas biaculeatus 10 Notes 11 References 12 External links

Distribution and habitats[edit] Anemonefish are native to warmer waters of the Indian and Pacific Oceans, including the Great Barrier Reef and the Red Sea. While most species have restricted distributions, others are widespread. Anemonefish live at the bottom of shallow seas in sheltered reefs or in shallow lagoons. No anemonefish are found in the Atlantic.[1]

Diet[edit] Anemonefish are omnivorous and can feed on undigested food from their host anemones, and the fecal matter from the anemonefish provides nutrients to the sea anemone. Anemonefish 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.[2][3] They may also consume the tentacles of their host anemone.[4]

Symbiosis and mutualism[edit] Anemonefish and sea anemones have a symbiotic, mutualistic relationship, each providing a number of benefits to the other. The individual species are generally highly host specific, and especially the genera Heteractis and Stichodactyla, and the species Entacmaea quadricolor are frequent anemonefish partners. The sea anemone protects the anemonefish from predators, as well as providing food through the scraps left from the anemone's meals and occasional dead anemone tentacles, and functions as a safe nest site. In return, the anemonefish defends the anemone from its predators and parasites.[5][6] The anemone also picks up nutrients from the anemonefish's excrement.[7] The nitrogen excreted from anemonefish increases the amount of algae incorporated into the tissue of their hosts, which aids the anemone in tissue growth and regeneration.[3] The activity of the anemonefish results in greater water circulation around the sea anemone,[8] and it has been suggested that their bright coloring might lure small fish to the anemone, which then catches them.[9] Studies on anemonefish have found that they alter the flow of water around sea anemone tentacles by certain behaviours and movements such as "wedging" and "switching". Aeration of the host anemone tentacles allows for benefits to the metabolism of both partners, mainly by increasing anemone body size and both anemonefish and anemone respiration.[10] Several theories are given about how they can survive the sea anemone poison: The mucus coating of the fish may be based on sugars rather than proteins. This would mean that anemones fail to recognize the fish as a potential food source and do not fire their nematocysts, or sting organelles. The coevolution of certain species of anemonefish with specific anemone host species may have allowed the fish to evolve an immunity to the nematocysts and toxins of their hosts. Amphiprion percula may develop resistance to the toxin from Heteractis magnifica, 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.[11] Anemonefish are the best known example of fish that are able to live among the venomous sea anemone tentacles, but several others occur, including juvenile threespot dascyllus, certain cardinalfish (such as Banggai cardinalfish), incognito (or anemone) goby, and juvenile painted greenling.[12][13][14]

Reproduction[edit] In a group of anemonefish, a strict dominance hierarchy exists. The largest and most aggressive female is found at the top. Only two anemonefish, a male and a female, in a group reproduce - through external fertilization. Anemonefish are sequential hermaphrodites, meaning they develop into males first, and when they mature, they become females. If the female anemonefish is removed from the group, such as by death, one of the largest and most dominant males becomes a female. The remaining males move up a rank in the hierarchy. Anemonefish lay eggs on any flat surface close to their host anemones. In the wild, anemonefish spawn around the time of the full moon. Depending on the species, they can lay hundreds or thousands of eggs. The male parent guards the eggs until they hatch about 6–10 days later, typically two hours after dusk.[15]

Parental investment[edit] Most anemonefish are protandrous hermaphrodites, meaning they alternate between the male and female sexes at some point in their lives. Anemonefish colonies usually consist of the reproductive male and female and a few juveniles, which help tend the colony.[16] Although multiple males cohabit an environment with a single female, polygamy does not occur and only the adult pair exhibits reproductive behavior. However, if the largest female dies, the social hierarchy shifts with the breeding male exhibiting protandrous sex reversal to become the breeding female. The largest juvenile then becomes the new breeding male after a period of rapid growth.[17] The existence of protandry in anemonefish may rest on the case that nonbreeders modulate their phenotype in a way that causes breeders to tolerate them. This strategy prevents conflict 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 fitness of the adult male, thereby protecting themselves from being evicted by the dominant fish.[18] The reproductive cycle of anemonefish is often correlated with the lunar cycle. Rates of spawning for anemonefish peak around the first and third quarters of the moon. The timing of this spawn means that the eggs 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 effectively sweeping them to safety. Before spawning, anemonefish exhibit increased rates of anemone and substrate biting, which help prepare and clean the nest for the spawn.[17] In terms of parental care, male anemonefish are often the caretakers of eggs. Before making the clutch, the parents often clear an oval-shaped clutch varying in diameter for the spawn. Fecundity, or reproductive rate, of the females usually ranges from 600 to 1500 eggs depending on her size. In contrast to most animal species, the female only occasionally takes responsibility for the eggs, with males expending most of the time and effort. Male anemonefish care for their eggs by fanning and guarding them for 6 to 10 days until they hatch. In general, eggs develop more rapidly in a clutch when males fan properly, and 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 different 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, though, display generally less preference for parental behavior than males. All these suggest that males have increased parental investment towards the eggs compared to females.[19]

In the aquarium[edit] Anemonefish make up 43% of the global marine ornamental trade, and 25% of the global trade comes from fish bred in captivity, while the majority is captured from the wild,[20][21] accounting for decreased densities in exploited areas.[22] Public aquaria and captive-breeding programs are essential to sustain their trade as marine ornamentals, and has recently become economically feasible.[23][24] It is one of a handful of marine ornamentals whose complete lifecycle has been closed in captivity. Members of some anemonefish species, such as the maroon clownfish, become aggressive in captivity; others, like the false percula clownfish, can be kept successfully with other individuals of the same species.[25] When a sea anemone is not available in an aquarium, the anemonefish may settle in some varieties of soft corals, or large polyp stony corals.[26] Once an anemone or coral has been adopted, the anemonefish will defend it. Anemonefish, however, are not obligately tied to hosts, and can survive alone in captivity.[27][28]

Taxonomy[edit] Historically, anemonefish have been identified by morphological features and color pattern in the field, while in a laboratory, other features such as scalation of the head, tooth shape, and body proportions are used.[2] These features have been used to group species into six complexes, clownfish, tomato, skunk, clarkii, saddleback, and maroon.[29] As can be seen from the gallery, each of the fish in these complexes has a similar appearance. Genetic analysis has shown that these complexes are not monophyletic groups, particularly the 11 species in the A. clarkii group, where only A. clarkii and A. tricintus are in the same clade, with six species,A. allardi A. bicinctus, A. chagosensis, A. chrosgaster, A. fuscocaudatus, A. latifasciatus, and A. omanensis being in an Indian clade, A. chrysopterus having monospecific lineage, and A. akindynos in the Australian clade with A. mccullochi.[30] Other significant differences are that A. latezonatus also has monospecific lineage, and A. nigripes is in the Indian clade rather than with A. akallopisos, the skunk anemonefish.[31] A. latezonatus is more closely related to A. percula and Premnas biaculeatus than to the saddleback fish with which it was previously grouped.[32][31] Obligate mutualism was thought to be the key innovation that allowed anemonefish to radiate rapidly, with rapid and convergent morhological changes correlated with the ecological niches offered by the host anemones.[32] The complexity of mitochondrial DNA structure shown by genetic analysis of the Australian clade suggested evolutionary connectivity among samples of A. akindynos and A. mccullochi that the authors theorize was the result of historical hybridization and introgression in the evolutionary past. The two evolutionary groups had individuals of both species detected, thus the species lacked reciprocal monophyly. No shared haplotypes were found between species.[33] Anemonefish Scientific name Common name Clade [30] Complex Genus Amphiprion:[34] Amphiprion akallopisos Skunk anemonefish A. akallopisos skunk A. akindynos Barrier reef anemonefish Australian A. clarkii A. allardi Allard's anemonefish Indian A. clarkii A. barberi Barber's anemonefish A. ephippium A. ephippium A. bicinctus Two-band anemonefish Indian A. clarkii A. chagosensis Chagos anemonefish Indian A. clarkii A. chrysogaster Mauritian anemonefish Indian A. clarkii A. chrysopterus Orange-fin anemonefish monospecific lineage A. clarkii A. clarkii Clark's anemonefish A. clarkii A. clarkii A. ephippium Red saddleback anemonefish A. ephippium A. ephippium A. frenatus Tomato anemonefish A. ephippium A. ephippium A. fuscocaudatus Seychelles anemonefish Indian [n 1] clarkii A. latezonatus Wide-band anemonefish monospecific lineage saddleback A. latifasciatus Madagascar anemonefish Indian A. clarkii A. leucokranos White-bonnet anemonefish likely hybrid skunk A. mccullochi Whitesnout anemonefish Australian A. ephippium A. melanopus Red and black anemonefish A. ephippium A. ephippium A. nigripes Maldive anemonefish Indian skunk A. ocellaris False clown anemonefish percula clownfish A. omanensis Oman anemonefish Indian A. clarkii A. pacificus Pacific anemonefish A. akallopisos skunk A. percula Clown anemonefish percula clownfish A. perideraion Pink skunk anemonefish A. akallopisos skunk A. polymnus Saddleback anemonefish A. polymnus saddleback A. rubrocinctus Australian anemonefish A. ephippium A. ephippium A. sandaracinos Orange anemonefish A. akallopisos skunk A. sebae Sebae anemonefish A. polymnus saddleback A. thiellei Thielle's anemonefish likely hybrid skunk A. tricinctus Three-band anemonefish clarkii clarkii Genus Premnas:[35] Premnas biaculeatus Maroon anemonefish percula Maroon

In popular culture[edit] This section needs expansion. You can help by adding to it. (November 2017) In Pixar's 2003 film Finding Nemo and its 2016 sequel Finding Dory, main characters Marlin and Nemo are clownfish.[36] The species depicted in the film is A. ocellaris.[5] The popularity of anenomefish greatly increased following the release of this film; greatly increasing the amount of captured specimens[37]

Gallery[edit] Clownfish[edit] A. ocellaris (false clown anemonefish) A. percula (clown anemonefish) in a 'normal' orange and a melanistic blackish variant A. clarkii complex[edit] A. clarkii (Clark's anemonefish) A. akindynos (Barrier Reef anemonefish) A. allardi (Allard's anemonefish). A. bicinctus (two-band anemonefish) A. chagosensis (Chagos anemonefish) A. chrysogaster (Mauritian anemonefish) A. chrysopterus (orange-fin anemonefish), Palau A. fuscocaudatus (Seychelles anemonefish) A. latifasciatus (Madagascar anemonefish) A. omanensis (Oman anemonefish) Tomato complex[edit] A. ephippium (red saddleback anemonefish) A. barberi (Barber's anemonefish) A. frenatus (tomato anemonefish) A. mccullochi (whitesnout anemonefish) A. melanopus (red and black anemonefish) A. rubrocinctus (Australian anemonefish) Saddleback complex[edit] A. latezonatus (wide-band anemonefish) A. polymnus (saddleback anemonefish) off Sulawesi, Indonesia A. sebae (sebae anemonefish) Skunk complex[edit] A. akallopisos (skunk anemonefish) A. leucokranos (white-bonnet anemonefish) A. nigripes (Maldive anemonefish) A. perideraion (pink skunk anemonefish) A. sandaracinos (orange anemonefish) A. perideraion (pink anemonefish) Yap, Micronesia Premnas biaculeatus[edit] P. biaculeatus (maroon anemonefish) P. biaculeatus (with yellow/gold striped bars)

Notes[edit] ^ Exemplars of A. fuscocaudatus have never been sequenced. The authors hypothetically placed this species in the Indian clade because it is the most parsimonious solution regarding the biogeography of anemonefish species.[30]

References[edit] ^ Society, National Geographic. "Clown Anemonefish, Clown Anemonefish Pictures, Clown Anemonefish Facts - National Geographic".  ^ a b Fautin, Daphne G.; Allen, Gerald R. (1997). Field Guide to Anemone Fishes and Their Host Sea Anemones. Western Australian Museum. ISBN 9780730983651. Archived from the original on 14 April 2015.  ^ a b Porat, D.; Chadwick-Furman, N.E. (2005). "Effects of anemonefish on giant sea anemones: Ammonium uptake, zooxanthella content and tissue regeneration" (PDF). Marine and Freshwater Behaviour and Physiology. 29 (1): 43–51. doi:10.1080/10236240500057929. Retrieved 18 April 2013.  ^ Fautin, D.G.; Guo, C.; Hwang, J.S. (1995). "Costs and benefits of the symbiosis between the anemoneshrimp Periclimenes brevicarpalis and its host Entacmaea quadricolor". Marine Ecology Progress Series. 129: 77–84. doi:10.3354/meps129077.  ^ a b "Clown Anemonefish". Nat Geo Wild : Animals. National Geographic Society. Retrieved 2011-12-19.  ^ Amphiprioninae at the Encyclopedia of Life ^ Holbrook, S. J. and Schmitt,R. J. Growth, reproduction and survival of a tropical sea anemone (Actiniaria): benefits of hosting anemonefish, 2005, cited in ^ Szczebak, Joseph T.; Raymond P. Henry; Fuad A. Al-Horani; Nanette E. Chadwick (2013). "Anemonefish oxygenate their anemone hosts at night". Journal of Experimental Biology. 216 (9): 970–976. doi:10.1242/jeb.075648. PMID 23447664.  ^ "Clown Anemonefishes, Amphiprion ocellaris". Marinebio. The MarineBio Conservation Society. Retrieved 2011-12-19.  ^ Joseph T. Szczebak; Raymond P. Henry; Fuad A. Al-Horani; Nanette E. Chadwick (2012-11-03). "Anemonefish oxygenate their anemone hosts at night". The Journal of Experimental Biology. Retrieved 2013-09-15.  ^ Mebs, D. 1994. "Anemonefish symbiosis: Vulnerability and Resistance of Fish to the Toxin of the Sea Anemone." Toxicon. Vol. 32(9):1059–1068. ^ Lieske, E.; and R. Myers (1999). Coral Reef Fishes. ISBN 0-691-00481-1 ^ Patzner, R.A. (5 July 2017). "Gobius incognitus". Retrieved 10 January 2018.  ^ Fretwell, K.; and B. Starzomski (2014). Painted greenling. Biodiversity of the Central Coast. Retrieved 29 January 2015. ^ Clownfish Breeding for Beginners by Jeff Hesketh of Mad Hatter's Reef ^ Stephanie Boyer. "Clown Anemofish". Florida Museum of Natural History. Retrieved 2013-09-15.  ^ a b Robert M. Ross (1978-02-10). "Reproductive Behavior of the Anemonefish Amphiprion melanopus on Guam" (PDF). Copeia. Archived from the original (PDF) on 2012-09-25. Retrieved 2013-09-15.  ^ Peter Buston (2004-08-18). "Does the presence of non-breeders enhance the fitness of breeders? An experimental analysis in the clown anemonefish Amphiprion percula" (PDF). Springer-Verlag. Retrieved 2013-09-15.  ^ Swagat Ghosh; T. T. Ajith Kumar; T. Balasubramanian (2011-08-04). "Determining the level of parental care relating fanning behavior of five species of clownfishes in captivity" (PDF). Indian Journal of Geo-Marine Sciences. Retrieved 2013-09-15.  ^ 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: 253–265. doi:10.1007/978-94-007-6016-5_17.  ^ Taylor, M.; Razak, T. & Green, E. (2003). From ocean to aquarium: A global trade in marine ornamental species (PDF). UNEP world conservation and monitoring centre (WCMC). pp. 1–64. Retrieved 18 April 2013.  ^ Shuman, Craig; Gregor Hodgson; Richard F. Ambrose (2005). "Population impacts of collecting sea anemones and anemonefish for the marine aquarium trade in the Philippines". Coral Reefs. 24: 564–573. doi:10.1007/s00338-005-0027-z.  ^ Watson, Craig; Jeffery Hill (2006). "Design criteria for recirculating, marine ornamental production systems". Aquacultural Engineering. 34 (3): 157–162. doi:10.1016/j.aquaeng.2005.07.002.  ^ Hall, Heather; Douglas Warmolts (2003). "23". In James C. Cato; Christopher L. Brown. Marine Ornamental Species: Collection, Culture and Conservation. Wiley-Blackwell. pp. 303–326. ISBN 978-0-8138-2987-6.  ^ Tullock, John (1998). Clownfish and Sea Anemones (illustrated ed.). Barron's Educational Series. pp. 11–22. Retrieved 2015-05-11.  ^ Fatherree, James W. "Aquarium Fish: On the Clownfishes' Range of Hosts". Advanced Aquarist. Retrieved 31 December 2016.  ^ Daphne Gail Fautin (1991). "The anemonefish symbiosis: what is known and what is not" (PDF). Symbiosis. 10: 23–46. Archived from the original (PDF) on 2012-05-25.  ^ Ronald L. Shimek (2004). Marine Invertebrates. Neptune City, NJ: T.F.H. Publications. p. 83. ISBN 978-1-890087-66-1.  ^ Goemans, B. "Anemonefishes". Retrieved 20 September 2015.  ^ a b c Litsios, G.; Salamin, N. (2014). "Hybridisation and diversification in the adaptive radiation of clownfishes". BMC Evolutionary Biology. 14: 245. doi:10.1186/s12862-014-0245-5. PMC 4264551 . PMID 25433367.  ^ a b DeAngelis, R. "What we really know about the diversity of Clownfish". Retrieved 20 September 2015.  ^ a b Litsios, G.; Sims, C.; Wüest, R.; Pearman P.B.; Zimmermann, N.E.; Salamin N. (2012). "Mutualism with sea anemones triggered the adaptive radiation of clownfishes". BMC Evolutionary Biology. 12 (11): 212. doi:10.1186/1471-2148-12-212. PMC 3532366 . PMID 23122007.  ^ van der Meer M.H.; G.P. Jones; J.-P.A. Hobbs; L. van Herwerden (2012). "Historic hybridization and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity". Ecology and Evolution. 2 (7): 1592–1604. doi:10.1002/ece3.251.  ^ Froese, Rainer, and Daniel Pauly, eds. (2011). Species of Amphiprion in FishBase. December 2011 version. ^ Froese, Rainer, and Daniel Pauly, eds. (2011). Species of Premnas in FishBase. December 2011 version. ^ "Finding Nemo (2003)". Retrieved 5 April 2016.  ^ Calado, Ricardo; Olivotto, Ike; Oliver, Miquel Planas; Holt, G. Joan (6 March 2017). "Marine Ornamental Species Aquaculture". John Wiley & Sons. p. 179 – via Google Books. 

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