Amphiprioninae

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Amphiprioninae
Amphiprion ocellaris (Clown anemonefish) by Nick Hobgood.jpg
Ocellaris clownfish, Amphiprion ocellaris
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Clade: Percomorpha
(unranked): Ovalentaria
Family: Pomacentridae
Subfamily: Amphiprioninae
Allen, 1975
Genera
Ocellaris clownfish nestled in a magnificent sea anemone (Heteractis magnifica) Amphiprion ocellaris (Clown anemonefish) in Heteractis magnifica (Sea anemone).jpg
Ocellaris clownfish nestled in a magnificent sea anemone (Heteractis magnifica)
A pair of pink anemonefish (Amphiprion perideraion) in their anemone home Skunk anemonefish.jpg
A pair of pink anemonefish ( Amphiprion perideraion ) in their anemone home
Clownfish swimming movements Nemo clown fish 1.gif
Clownfish swimming movements
A clownfish swimming Clown Fish Swimming.jpg
A clownfish swimming
Video of a cinnamon clownfish swimming around an anemone

Clownfish or anemonefish are fishes 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, anemonefish are overall yellow, orange, or a reddish or blackish color, and many show white bars or patches. The largest can reach a length of 17 cm (6+1⁄2 in), while the smallest barely achieve 7–8 cm (2+3⁄4–3+1⁄4 in).

Contents

Distribution and habitats

Anemonefish are endemic to the warmer waters of the Indian Ocean, including the Red Sea, and Pacific Ocean, the Great Barrier Reef, Southeast Asia, Japan, and the Indo-Malaysian region. While most species have restricted distributions, others are widespread. Anemonefish typically live at the bottom of shallow seas in sheltered reefs or in shallow lagoons. No anemonefish are found in the Atlantic. [1]

Diet

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]

Symbiosis and mutualism

Anemonefish and sea anemones have a symbiotic, mutualistic relationship, each providing many 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. [4] [5] The anemone also picks up nutrients from the anemonefish's excrement. [6] The nitrogen excreted from anemonefish increases the number 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, [7] and it has been suggested that their bright coloring might lure small fish to the anemone, which then catches them. [8] Studies on anemonefish have found that they alter the flow of water around sea anemone tentacles by certain behaviors 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. [9]

Bleaching of the host anemone can occur when warm temperatures cause a reduction in algal symbionts within the anemone. Bleaching of the host can cause a short-term increase in the metabolic rate of resident anemonefish, probably as a result of acute stress. [10] Over time, however, there appears to be a down-regulation of metabolism and a reduced growth rate for fish associated with bleached anemones. These effects may stem from reduced food availability (e.g. anemone waste products, symbiotic algae) for the anemonefish. [11]

Several theories are given about how they can survive the sea anemone poison:

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. [13] [14] [15]

Reproduction

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 protandrous 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. [16]

Parental investment

Anemonefish colonies usually consist of the reproductive male and female and a few male juveniles, which help tend the colony. [17] 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 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. [18] 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 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. [19]

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 protect 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. [18]

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 can 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. [20]

Taxonomy

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, percula, tomato, skunk, clarkii, saddleback, and maroon. [21] 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. allardiA. 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. [22] 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. [23] A. latezonatus is more closely related to A. percula and Premnas biaculeatus than to the saddleback fish with which it was previously grouped. [24] [23]

Obligate mutualism was thought to be the key innovation that allowed anemonefish to radiate rapidly, with rapid and convergent morphological changes correlated with the ecological niches offered by the host anemones. [24] 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. [25]

Phylogenetic relationships

Scientific nameCommon name Clade [22] Complex
Genus Amphiprion: [26]
Amphiprion akallopisos Skunk anemonefishA. akallopisosSkunk
A. akindynos Barrier reef anemonefishAustralianA. clarkii
A. allardi Allard's anemonefishIndianA. clarkii
A. barberi Barber's anemonefishA. ephippium A. ephippium
A. bicinctus Two-band anemonefishIndianA. clarkii
A. chagosensis Chagos anemonefishIndianA. clarkii
A. chrysogaster Mauritian anemonefishIndianA. clarkii
A. chrysopterus Orange-fin anemonefish Monospecific lineageA. clarkii
A. clarkii Clark's anemonefishA. clarkiiA. clarkii
A. ephippium Red saddleback anemonefishA. ephippiumA. ephippium
A. frenatus Tomato anemonefishA. ephippiumA. ephippium
A. fuscocaudatus Seychelles anemonefishIndian [n 1] Clarkii
A. latezonatus Wide-band anemonefish Monospecific lineageSaddleback
A. latifasciatus Madagascar anemonefishIndianA. clarkii
A. leucokranos White-bonnet anemonefishLikely hybridSkunk
A. mccullochi Whitesnout anemonefishAustralianA. ephippium
A. melanopus Red and black anemonefishA. ephippium A. ephippium
A. nigripes Maldive anemonefishIndianSkunk
A. ocellaris False clown anemonefishPerculaClownfish
A. omanensis Oman anemonefishIndianA. clarkii
A. pacificus Pacific anemonefishA. akallopisosSkunk
A. percula Clown anemonefishPerculaClownfish
A. perideraion Pink skunk anemonefishA. akallopisosSkunk
A. polymnus Saddleback anemonefishA. polymnusSaddleback
A. rubrocinctus Australian anemonefishA. ephippiumA. ephippium
A. sandaracinos Orange anemonefishA. akallopisosSkunk
A. sebae Sebae anemonefishA. polymnusSaddleback
A. thiellei Thielle's anemonefishLikely hybridSkunk
A. tricinctus Three-band anemonefishClarkiiClarkii
Genus Premnas: [27]
Premnas biaculeatus Maroon anemonefishPerculaMaroon

Morphological diversity by complex

In the aquarium

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, [28] [29] accounting for decreased densities in exploited areas. [30] Public aquaria and captive-breeding programs are essential to sustain their trade as marine ornamentals, and has recently become economically feasible. [31] [32] It is one of a handful of marine ornamentals whose complete lifecycle has been in closed 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. [33]

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. [34] 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. [35] [36]

In Disney Pixar's 2003 film Finding Nemo and its 2016 sequel Finding Dory main characters Nemo, his father Marlin, and his mother Coral are clownfish, probably the species A. ocellaris . [37] The popularity of anemonefish for aquaria increased following the film's release; it is the first film associated with an increase in the numbers of those captured in the wild. [38]

Notes

  1. ↑ 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. [22]

Related Research Articles

<span class="mw-page-title-main">Orange clownfish</span> Species of fish

The orange clownfish also known as percula clownfish and clown anemonefish, is widely known as a popular aquarium fish. Like other clownfishes, it often lives in association with sea anemones. A. percula is associated specifically with Heteractis magnifica and Stichodactyla gigantea, and as larvae use chemical cues released from the anemones to identify and locate the appropriate host species to use them for shelter and protection. This causes preferential selection when finding their anemone host species. Although popular, maintaining this species in captivity is rather complex. The Great Barrier Reef Marine Park Authority regulates the number of collection permits issued to aquarium fish dealers who seek this, and other tropical fish within the Great Barrier Reef Marine Park. The symbiosis between anemonefish and anemones depends on the presence of the fish drawing other fish to the anemone, where they are stung by its venomous tentacles. The anemone helps the fish by giving it protection from predators, which include brittle stars, wrasses, and other damselfish, and the fish helps the anemone by feeding it, increasing oxygenation, and removing waste material from the host. Various hypotheses exist about the fish's ability to live within the anemone without being harmed. One study carried out at Marineland of the Pacific by Dr. Demorest Davenport and Dr. Kenneth Noris in 1958 revealed that the mucus secreted by the anemone fish prevented the anemone from discharging its lethal stinging nematocysts. A second hypothesis is that A. percula has acquired immunity towards the sea anemone's toxins, and a combination of the two has been shown to be the case. The fish feed on algae, zooplankton, worms, and small crustaceans.

<span class="mw-page-title-main">Clark's anemonefish</span> Species of fish

Amphiprion clarkii, known commonly as Clark's anemonefish and yellowtail clownfish, is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes.

<span class="mw-page-title-main">Ocellaris clownfish</span> Species of fish

The ocellaris clownfish, also known as the false percula clownfish or common clownfish, is a marine fish belonging to the family Pomacentridae, which includes clownfishes and damselfishes. Amphiprion ocellaris are found in different colors, depending on where they are located. For example, black Amphiprion ocellaris with white bands can be found near northern Australia, Southeast Asia, and Japan. Orange or red-brown Amphiprion ocellaris also exist with three similar white bands on the body and head. Amphiprion ocellaris can be distinguished from other Amphiprion species based on the number of pectoral rays and dorsal spines. Amphiprion ocellaris are known to grow about 110 mm long. Like many other fish species, females are, however, larger than males. The life cycle of Amphiprion ocellaris varies in whether they reside at the surface or bottom of the ocean. When they initially hatch, they reside near the surface. However, when Amphiprion ocellaris enter into the juvenile stage of life, they travel down to the bottom to find shelter in a host anemone. Once they find their anemone, they form a symbiotic relationship with them.

<span class="mw-page-title-main">Tomato clownfish</span> Species of fish

The tomato clownfish is a species of marine fish in the family Pomacentridae, the clownfishes and damselfishes. It is native to the waters of the Western Pacific, from the Japan to Indonesia. Other common names include blackback anemonefish, bridled anemonefish, fire clown, and red tomato clown.

<span class="mw-page-title-main">Orange skunk clownfish</span> Species of fish

Amphiprion sandaracinos, also known as the orange skunk clownfish or orange anemonefish, is a species of anemonefish that is distinguished by its broad white stripe along the dorsal ridge. Like all anemonefishes it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of the host anemone. It is a sequential hermaphrodite with a strict sized based dominance hierarchy: the female is largest, the breeding male is second largest, and the male non-breeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male will change to female if the sole breeding female dies, with the largest non-breeder becomes the breeding male.

<span class="mw-page-title-main">Sebae clownfish</span> Species of fish

Amphiprion sebae, also known as the sebae clownfish, is an anemonefish found in the northern Indian Ocean, from Java to the Arabian Peninsula. Like all anemonefish it is usually found living in association with sea anemones. While the common name of Heteractis crispa, the sebae anemone, suggests an association, it is normally found with the Stichodactyla haddoni or saddle anemone. A. sebae, like all anemonefish, lives in a symbiotic relationship with the host anemone where the fish is unaffected by the stinging tentacles of the anemone. In a group of clownfish, Only two clownfish, a male and a female, in a group reproduce through external fertilization. Clownfish are sequential hermaphrodites, changing from male to female, with a strict dominance hierarchy and only the largest fish being female.

<span class="mw-page-title-main">Sebae anemone</span> Species of sea anemone

The sebae anemone, also known as leathery sea anemone, long tentacle anemone, or purple tip anemone, is a species of sea anemone belonging to the family Stichodactylidae and native to the Indo-Pacific area.

<span class="mw-page-title-main">Orange-fin anemonefish</span> Species of fish

The orange-fin anemonefish is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes, found in the Western Pacific north of the Great Barrier Reef from the surface to 20 m, to include the Pacific Ocean between Queensland, Australia, and New Guinea to the Marshall and Tuamotus Islands. It can grow to 17 cm in length.

<span class="mw-page-title-main">Allard's clownfish</span> Species of fish

Allard's clownfish or Allard's anemonefish is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes, from the western Indian Ocean off the coast of East Africa and the Mascarenes.

<span class="mw-page-title-main">Saddleback clownfish</span> Species of fish

Amphiprion polymnus, also known as the saddleback clownfish or yellowfin anemonefish, is a black and white species of anemonefish with a distinctive saddle. Like all anemonefishes it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of the host anemone. It is a sequential hermaphrodite with a strict sized-based dominance hierarchy: the female is largest, the breeding male is second largest, and the male non-breeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male will change to female if the sole breeding female dies, with the largest non-breeder becomes the breeding male.

<i>Amphiprion akindynos</i> Species of fish

Amphiprion akindynos, the Barrier Reef anemonefish, is a species of anemonefish that is principally found in the Great Barrier Reef of Australia, but also in nearby locations in the Western Pacific. The species name 'akindynos' is Greek, meaning 'safe' or 'without danger' in reference to the safety afforded amongst the tentacles of its host anemone. Like all anemonefishes it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of the host anemone. It is a sequential hermaphrodite with a strict size-based dominance hierarchy: the female is largest, the breeding male is second largest, and the male non-breeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male will change to female if the sole breeding female dies, with the largest non-breeder becomes the breeding male. The fish's natural diet includes zooplankton.

<span class="mw-page-title-main">Red Sea clownfish</span> Species of fish

Amphiprion bicinctus, meaning "both sawlike with two stripes," commonly known as the Red Sea or two-bandedanemonefish is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes. Like other species of the genus, the fish feeds on algae and zooplankton in the wild.

<span class="mw-page-title-main">Pink skunk clownfish</span> Species of fish

Amphiprion perideraion, also known as the pink skunk clownfish or the pink anemonefish, is a species of anemonefish that is widespread from northern Australia through the Malay Archipelago and Melanesia. Like all anemonefishes, it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of the host. It is a sequential hermaphrodite with a strict size-based dominance hierarchy; the female is largest, the breeding male is second largest, and the male nonbreeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male changes to female if the sole breeding female dies, with the largest nonbreeder becoming the breeding male.

<span class="mw-page-title-main">Red saddleback anemonefish</span> Species of fish

The red saddleback anemonefish, Amphiprion ephippium, also known as the saddle anemonefish, is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes.

The three-band anemonefish is a species of anemonefish endemic to the Marshall Islands in the western part of the Pacific Ocean. Like all anemonefishes, it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of its host. It is a sequential hermaphrodite with a strict size-based dominance hierarchy; the female is largest, the breeding male is second largest, and the male nonbreeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male changes to female if the sole breeding female dies, with the largest nonbreeder becoming the breeding male. The fish's natural diet includes zooplankton.

<i>Stichodactyla gigantea</i> Species of sea anemone

Stichodactyla gigantea, commonly known as the giant carpet anemone, is a species of sea anemone that lives in the Indo-Pacific area. It can be kept in an aquarium but is a very challenging species to keep alive and healthy for more than 3–5 years.

<i>Amphiprion latezonatus</i> Species of fish

Amphiprion latezonatus, also known as the wide-band anemonefish, is a species of anemonefish found in subtropical waters off the east coast of Australia. Like all anemonefishes, it forms a symbiotic mutualism with sea anemones and is unaffected by the stinging tentacles of its host. It is a sequential hermaphrodite with a strict size-based dominance hierarchy; the female is largest, the breeding male is second largest, and the male nonbreeders get progressively smaller as the hierarchy descends. They exhibit protandry, meaning the breeding male changes to female if the sole breeding female dies, with the largest nonbreeder becoming the breeding male.

<i>Amphiprion chagosensis</i> Species of fish

Amphiprion chagosensis, the Chagos anemonefish, is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes. It is named for the Chagos Archipelago in the Indian Ocean and it is endemic to the archipelago. The original specimens were collected at Diego Garcia Atoll, Chagos Archipelago.

<i>Amphiprion fuscocaudatus</i> Species of fish

Amphiprion fuscocaudatus is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes.

<i>Amphiprion latifasciatus</i> Species of fish

Amphiprion latifasciatus is a marine fish belonging to the family Pomacentridae, the clownfishes and damselfishes.

References

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  2. 1 2 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.
  3. 1 2 Porat, D.; Chadwick-Furman, N.E. (March 2005). "Effects of anemonefish on giant sea anemones: Ammonium uptake, zooxanthella content and tissue regeneration". Marine and Freshwater Behaviour and Physiology. 38 (1): 43–51. doi:10.1080/10236240500057929. S2CID   53051081.
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  10. ↑ Norin, Tommy; Mills, Suzanne; Crespel, Amelie; Cortese, Daphne; Beldade, Ricardo; Killen, Shaun (2018). "Anemone bleaching increases the metabolic demands of symbiont anemonefish". Proceedings of the Royal Society B. 285 (1876). doi: 10.1098/rspb.2018.0282 . PMC   5904320 . PMID   29643214.
  11. ↑ Cortese, Daphne; Norin, Tommy; Beldade, Ricardo; Crespel, Amelie; Killen, Shaun; Mills, Suzanne (2021). "Physiological and behavioural effects of anemone bleaching on symbiont anemonefish in the wild". Functional Ecology. 35 (3): 663–674. doi: 10.1111/1365-2435.13729 .
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  15. ↑ Fretwell, K.; and B. Starzomski (2014). Painted greenling. Biodiversity of the Central Coast. Retrieved 29 January 2015.
  16. ↑ Clownfish Breeding for Beginners by Jeff Hesketh of Mad Hatter's Reef
  17. ↑ Stephanie Boyer. "Clown Anemofish". Florida Museum of Natural History. Archived from the original on October 28, 2005. Retrieved 2013-09-15.
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  22. 1 2 3 Litsios, Glenn; Salamin, Nicolas (December 2014). "Hybridisation and diversification in the adaptive radiation of clownfishes". BMC Evolutionary Biology. 14 (1): 245. doi:10.1186/s12862-014-0245-5. PMC   4264551 . PMID   25433367.
  23. 1 2 DeAngelis, R. "What we really know about the diversity of Clownfish". Archived from the original on 2015-09-25. Retrieved 20 September 2015.
  24. 1 2 Litsios, Glenn; Sims, Carrie A; Wüest, Rafael O; Pearman, Peter B; Zimmermann, Niklaus E; Salamin, Nicolas (2012). "Mutualism with sea anemones triggered the adaptive radiation of clownfishes". BMC Evolutionary Biology. 12 (1): 212. doi:10.1186/1471-2148-12-212. PMC   3532366 . PMID   23122007.
  25. ↑ van der Meer, M. H.; Jones, G. P.; Hobbs, J.-P. A.; van Herwerden, L. (July 2012). "Historic hybridization and introgression between two iconic Australian anemonefish and contemporary patterns of population connectivity: Historic Hybridization between Anemonefish". Ecology and Evolution. 2 (7): 1592–1604. doi:10.1002/ece3.251. PMC   3434915 . PMID   22957165.
  26. ↑ Froese, Rainer and Pauly, Daniel, eds. (2011). Species of Amphiprion in FishBase . December 2011 version.
  27. ↑ Froese, Rainer and Pauly, Daniel, eds. (2011). Species of Premnas in FishBase . December 2011 version.
  28. ↑ Dhaneesh, K. V.; Vinoth, R.; Ghosh, Swagat; Gopi, M.; Kumar, T. T. Ajith; Balasubramanian, T. (2013). "Hatchery Production of Marine Ornamental Fishes: An Alternate Livelihood Option for the Island Community at Lakshadweep". Climate Change and Island and Coastal Vulnerability. pp. 253–265. doi:10.1007/978-94-007-6016-5_17. ISBN   978-94-007-6015-8.
  29. ↑ 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. Archived from the original (PDF) on July 1, 2004. Retrieved 18 April 2013.
  30. ↑ Shuman, Craig S.; Hodgson, Gregor; Ambrose, Richard F. (December 2005). "Population impacts of collecting sea anemones and anemonefish for the marine aquarium trade in the Philippines". Coral Reefs. 24 (4): 564–573. Bibcode:2005CorRe..24..564S. doi:10.1007/s00338-005-0027-z. S2CID   25027153.
  31. ↑ Watson, Craig A.; Hill, Jeffrey E. (May 2006). "Design criteria for recirculating, marine ornamental production systems". Aquacultural Engineering. 34 (3): 157–162. doi:10.1016/j.aquaeng.2005.07.002.
  32. ↑ Hall, Heather; Douglas Warmolts (2003). "23". In James C. Cato; Christopher L. Brown (eds.). Marine Ornamental Species: Collection, Culture and Conservation. Wiley-Blackwell. pp. 303–326. ISBN   978-0-8138-2987-6.
  33. ↑ Tullock, John (1998). Clownfish and Sea Anemones (illustrated ed.). Barron's Educational Series. pp. 11–22. ISBN   9780764105111 . Retrieved 2015-05-11.
  34. ↑ Fatherree, James W. "Aquarium Fish: On the Clownfishes' Range of Hosts". Advanced Aquarist. Archived from the original on March 22, 2014. Retrieved 31 December 2016.
  35. ↑ 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.
  36. ↑ Ronald L. Shimek (2004). Marine Invertebrates. Neptune City, NJ: T.F.H. Publications. p. 83. ISBN   978-1-890087-66-1.
  37. ↑ "Finding Nemo (2003)". Rotten Tomatoes. Retrieved 5 April 2016.
  38. ↑ Calado, Ricardo; Olivotto, Ike; Oliver, Miquel Planas; Holt, G. Joan (6 March 2017). Marine Ornamental Species Aquaculture. John Wiley & Sons. p. 179. ISBN   9780470673904 – via Google Books.

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