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Conservation Status ( anglais )

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Queen angelfish are considered a species of least concern by the ICUN. Populations are globally stable, although they are harvested in high numbers near Brazil.

US Federal List: no special status

CITES: no special status

IUCN Red List of Threatened Species: least concern

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Associations ( anglais )

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Queen angelfish may be preyed upon by many larger fish that inhabit coral reefs; however, predation has not been well studied.

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Morphology ( anglais )

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Queen angelfish are easily distinguished by their striking coloration, with vibrant yellow accents and variations of gem-like blues. Their distinctive "crown" is speckled dark blue and surrounded by a ring of bright blue. Their tail is yellow. Juveniles have a markedly different coloration than adults, displaying a striped blue and yellow pattern or a solid yellow pattern.

The body of queen angelfish is very flat, with an elongated, continuous dorsal and anal fin with 9 to 15 spines and 15 to 17 soft rays. They have a strong spine at the angle of the preopercle (cheek bone) and lack a well developed pelvic axillary process (fleshy bump at the base of the pelvic fin). Queen angelfish average 45 cm in length and 1.6 kg in mass. Males are generally larger than females.

Average mass: 1.6 kg.

Average length: 45 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: male larger

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Life Expectancy ( anglais )

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The lifespan of queen angelfish has not been well documented.

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Habitat ( anglais )

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Queen angelfish are primarily found in coral reefs, which provide shelter and abundant food sources. They can be found at depths up to 70 m. Although they are naturally marine fish, queen angelfish can tolerate changes in salinity. As such, they are often placed in marine aquariums.

Range depth: 70 (high) m.

Habitat Regions: tropical ; saltwater or marine

Aquatic Biomes: reef

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Distribution ( anglais )

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Queen angelfish, Holacanthus ciliaris, are tropical fish found in coral reefs in the western Pacific Ocean, the Indian Ocean, and the Atlantic Ocean as far south as Brazil. They do not migrate, and they are commonly spotted near the Bahamas and Florida.

Biogeographic Regions: indian ocean (Native ); atlantic ocean (Native ); pacific ocean (Native )

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Trophic Strategy ( anglais )

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Queen angelfish primarily feed on sponges and corals. They also eat other marine invertebrates, including tunicates, jellyfish, hydroids, bryozoans. They may also eat plankton and algae.

Animal Foods: cnidarians; other marine invertebrates; zooplankton

Plant Foods: algae

Primary Diet: carnivore (Eats other marine invertebrates)

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Associations ( anglais )

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Queen angelfishes feed on sponges, corals and other small invertebrates, and are preyed upon by larger animals that inhabit coral reefs.

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Benefits ( anglais )

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Queen angelfish are popular additions to saltwater aquariums because of their beautiful coloration as both juveniles and adults. Because new technologies have allowed hobbyists to effectively care for and keep marine fish, queen angelfish are increasing sought after. In Florida, queen angelfish averaged from $11.16 to $17.84 USD per fish between 1990 and 1998. Retail prices vary with size and range between $60 and $130 USD. Adult mating pairs sell at a premium.

Positive Impacts: pet trade

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Benefits ( anglais )

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There are no known adverse effects of queen angelfish on humans.

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Life Cycle ( anglais )

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After fertilized eggs of queen angelfish float in the water column for 15 to 20 hours, they develop into transparent larvae. Larvae then develop into juveniles, which resemble adults. Larvae feed on plankton and grow rapidly, reaching a size of 15 to 20 mm in their juvenile form.

Development - Life Cycle: metamorphosis

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Behavior ( anglais )

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Queen angelfish communicate, particularly during mating, through temporary changes in color. Little information is otherwise available regarding the communication and perception of queen angelfish.

Communication Channels: visual

Perception Channels: visual ; tactile ; chemical

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Reproduction ( anglais )

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Queen angelfish are believed to be polygynous, and harems have been observed during courtship and pre-spawning. Harems generally consist of 1 male and up to 4 females. A male courts a female by displaying his pectoral fins, flicking them outward every few seconds. The female then ascends in the water, and the male positions himself below the female. The male touches his snout to her vent (genital) area, rising with the female with his belly close to hers. As the pair rises to about 18 m in depth, they release eggs and sperm.

Mating System: polygynous

Queen angelfish spawn seasonally, which occurs during the winter months in Puerto Rico. Spawning peaks once each year, although queen angelfish may spawn more than once during the year. Spawning behavior has been observed within minutes of sunset during the evening. Females can produce 25,000 to 75,000 eggs in one evening. Eggs hatch in 15 to 20 hours, and larvae absorb the yolk sac in the next 48 hours. Larvae feed on plankton and grow rapidly, reaching a size of 15 to 20 mm in their juvenile form.

Breeding interval: Spawning peaks once a year, but queen angelfish may spawn more than once during the year.

Breeding season: Queen angelfish spawn seasonally, which occurs during the winter in Puerto Rico.

Range gestation period: 15 to 20 hours.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (External ); broadcast (group) spawning; oviparous

Once eggs are fertilized, zygotes are left develop into larvae without any parental investment. Juvenile queen angelfish find protection among colonies of finger sponges and corals at the bottom of reefs.

Parental Investment: no parental involvement; pre-fertilization (Provisioning)

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Ajagbe, O. 2011. "Holacanthus ciliaris" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Holacanthus_ciliaris.html
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O. Omodele Ajagbe, University of Michigan-Ann Arbor
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Phil Myers, University of Michigan-Ann Arbor
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Gail McCormick, Special Projects
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Diagnostic Description ( anglais )

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Tail and pectoral fins entirely yellow. Black spot on forehead has electric blue spots and is surrounded by narrow, electric blue ring. Large blue spot at base of pectoral fin (Ref. 26938). Adults develop short spines on the margin. The color of large adults is purplish blue with yellow-orange rims to the scales; head above eye dark blue, below greenish yellow; mouth, chin, throat, chest and abdomen purplish blue (Ref. 13442)
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Morphology ( anglais )

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Dorsal spines (total): 14; Dorsal soft rays (total): 19 - 21; Analspines: 3; Analsoft rays: 20 - 21
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Trophic Strategy ( anglais )

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Found on coral reefs. Moves gracefully between seafans, seawhips, and corals (Ref. 9710). Stomach contents of 26 specimens indicate that the species feeds almost exclusively on sponges supplemented by small amounts of algae, tunicates, hydroids and bryozoans. Omnivore (Ref. 57616).
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Biology ( anglais )

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Found on coral reefs. Generally occurs solitarily or in pairs. Moves gracefully between seafans, seawhips, and corals (Ref. 9710). Stomach contents of 26 specimens indicate that the species feeds almost exclusively on sponges supplemented by small amounts of algae, tunicates, hydroids and bryozoans. Young pick ectoparasites from other fishes (Ref. 9710). Marketed fresh (Ref. 3797).
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Importance ( anglais )

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fisheries: minor commercial; aquarium: commercial; price category: high; price reliability: very questionable: based on ex-vessel price for species in this family
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Queen angelfish ( anglais )

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The queen angelfish (Holacanthus ciliaris), also known as the blue angelfish, golden angelfish, or yellow angelfish, is a species of marine angelfish found in the western Atlantic Ocean. It is a benthic (ocean floor) warm-water species that lives in coral reefs. It is recognized by its blue and yellow coloration and a distinctive spot or "crown" on its forehead. This crown distinguishes it from the closely related and similar-looking Bermuda blue angelfish (Holacanthus bermudensis), with which it overlaps in range and can interbreed.

Adult queen angelfish are selective feeders and primarily eat sponges. Their social structure consists of harems which include one male and up to four females. They live within a territory where the females forage separately and are tended to by the male. Breeding in the species occurs near a full moon. The transparent eggs float in the water until they hatch. Juveniles of the species have different coloration than adults and act as cleaner fish.

The queen angelfish is popular in the aquarium trade and has been a particularly common exported species from Brazil. In 2010, the queen angelfish was assessed as least concern by the International Union for Conservation of Nature as the wild population appeared to be stable.

Taxonomy

The queen angelfish was first described as Chaetodon ciliaris in 1758 by Carl Linnaeus in the 10th edition of his Systema Naturae, with the type locality given as the "Western Atlantic/Caribbean".[3] In 1802 it was moved by French naturalist Bernard Germain de Lacépède to the genus Holacanthus,[4] the name of which is derived from the Ancient Greek words "holos" (full) and "akantha" (thorn).[2] Its specific name ciliaris means "fringed", a reference to its squamis ciliatis ("ciliate scales").[5] Other common names for the species include "blue angelfish", "golden angelfish" and "yellow angelfish".[6]

Marine angelfish of the genus Holacanthus likely emerged between 10.2 and 7.6 million years ago (mya). The most basal species is the Guinean angelfish (Holacanthus africanus) off the coast of West Africa, indicating that the lineage colonized the Atlantic from the Indian Ocean.[7] The closure of the Isthmus of Panama 3.5–3.1 mya led to the splitting off of the Tropical Eastern Pacific species.[8] The closest relative and sister species of the queen angelfish is the sympatric and similar Bermuda blue angelfish (H. bermudensis), from which it split around 1.5 mya.[7][8] They are known to interbreed, producing a hybrid known as the Townsend angelfish[9] which has features similar to both parent species.[10] The Townsend angelfish is fertile, and individuals can breed both with each other and with the two parent species.[11]

The following cladogram is based on molecular evidence:[7][8][12]

Image of a fish in a coral reef
Townsend angelfish, a hybrid of queen and Bermuda blue angelfish
Holacanthus

Rock beauty Holacanthus tricolor 1 (cropped).jpg

Bermuda blue angelfish Bermuda blue angelfish (cropped).jpg

Queen angelfish Holacanthus ciliaris 10 (cropped).jpg

Clarion angelfish Clarion angelfish (Holacanthus clarionensis) (19185438555) (cropped).jpg

Clipperton angelfish Holacanthus limbaughi.jpg

King angelfish Holacanthus passer 1.jpg

Guinean angelfish Holacanthus africanus.jpg

Description

Frontal view of a queen angelfish showing the crown
With "crown" visible

The queen angelfish has a broad, flattened, oval-shaped body with a reduced, dulled snout and a small mouth containing bristle-like teeth.[13] The dorsal fin contains 14 spines and 19–21 soft rays, and the anal fin has 3 spines and 20–21 soft rays.[2] This species attains a maximum total length of 45 cm (18 in) and weight of 1,600 g (56 oz).[2][6] Males may be larger than females.[14][15]

The species is covered in yellow-tipped blue-green scales, with a bright yellow tail, pectoral and pelvic fins. Both the dorsal and anal fins have orange-yellow end points, while the pectoral fins have blue patches at the base.[14] On the forehead is an eye-like spot or "crown" that is cobalt blue with an electric blue outer ring and dotted with electric blue spots.[2][6][14] This crown is the main feature distinguishing the species from the Bermuda blue angelfish. Juveniles are dark blue with bright blue vertical stripes and a yellow pectoral area. They resemble juvenile blue angelfish and are distinguished by more curved vertical stripes.[6] Growing juveniles develop transitional patterns as they reach their adult coloration.[14]

Juvenile closer to adult coloration

Seven other color morphs have been recorded off the coast of the Saint Peter and Saint Paul Archipelago, Brazil. The most commonly recorded is a mostly gold or bright orange morph. Other morphs may be bright blue with some yellow, black or white coloration or even all white.[16] Another color morph was recorded off Dry Tortugas, Florida, in 2009. This fish was mostly cobalt blue with white and yellow-orange colored areas.[17]

There are records of at least two wild queen angelfish at St. Peter and St. Paul with a "pughead" skeletal deformity, a squashed upper jaw and a lower jaw that sticks out. Such deformities mostly occur in captive fish.[18]

Ecology

Queen angelfish are found in tropical and subtropical areas of the Western Atlantic Ocean around the coasts and islands of the Americas. They occur from Florida along the Gulf of Mexico and the Caribbean Sea down to Brazil. Their range extends as far east as Bermuda and the Saint Peter and Saint Paul Archipelago.[1][6] Queen angelfish are benthic or bottom-dwelling and occur from shallow waters close to shore down to 70 m (230 ft). They live in coral reefs, preferring soft corals, and swim either alone or in pairs.[6]

Queen angelfish eat sponges, tunicates, jellyfish, corals, plankton, and algae. Juveniles act as cleaner fish and establish and remove ectoparasites from bigger fish.[6] Off St. Thomas Island and Salvador, Bahia, 90% of the diet of adults is sponges. Off the Saint Peter and Saint Paul Archipelago, more than 30 prey species may be consumed, 68% being sponges, 25% being algae, and 5% being bryozoans. Queen angelfish appear to be selective feeders as the proportion of prey in their diet does not correlate with their abundance. On the species level, the angelfish of the Saint Peter and Saint Paul Archipelago target the less common sponges Geodia neptuni, Erylus latens, Clathria calla, and Asteropus niger.[19]

Life cycle

Male queen angelfish have large territories with a harem of two to four females.[15] Little is known about the sexual development of the species, though they are presumed to be protogynous hermaphrodites. The largest harem female may transform into a male if the territorial male disappears.[11] Around midday, the females forage individually in different locations. The male tends to each of them, rushing at, circling, and feeding next to them.[15] Spawning in this species occurs year-round.[20] It is observed sometime around a full moon.[15]

Courtship involves the male showing his side to the female and flick his pectoral fins at her or "soaring" above them. At the beginning of spawning, the female swims towards the surface with the male swimming under her with his snout pressing against her vent.[15] They then deposit their eggs and semen into the water. The female discharges between 25 and 75 thousand eggs a day.[6] After spawning, the pair split and head to back to the ocean floor.[11]

The transparent eggs are pelagic and remain suspended in the water for 15–20 hours. The hatched larvae have a large yolk sac with no functional eyes, gut or fins, but two days later, the yolk is absorbed, and the larvae have more of a resemblance to fish. These larvae are plankton-eaters and grow quickly. Between the ages of three and four weeks old, when they have reached a length of 15 to 20 mm (0.6 to 0.8 in), they descend to the floor as juveniles. Juvenile angelfish live alone and in territories encompassing finger sponges and coral, where they establish cleaning stations for other fish.[6]

Human interactions

Aquarium in Barcelona, Spain

Queen angelfish are not normally eaten or commercially fished. They are captured mostly for the aquarium trade, where they are highly valued.[21] As juveniles, angelfish can adapt to eating typical aquarium food and hence have a higher survival rate than individuals taken as adults, which require a more specialized diet.[22]

In Brazil, the queen angelfish is the most common marine ornamental fish sold aboard.[1] From 1995 to 2000, 43,730 queen angelfish were traded at Fortaleza in the northeast of the country, and in 1995, 75% of marine fish sold were both queen and French angelfish.[23] In 2010, the queen angelfish was assessed as least concern by the International Union for Conservation of Nature, as the species is only significantly fished off Brazil and the wild population appeared to be secure.[1]

Queen angelfish have been caught in the eastern Adriatic Sea, off Croatia, in 2011, and the Mediterranean Sea, off Malta, in 2020. These are likely introductions from the aquarium industry and not natural colonizations.[24] In 2015, an aquarium-introduced angelfish was found in the Red Sea at Eilat's Coral Beach, Israel. Its kidney was infected with the disease-causing bacterium Photobacterium damselae piscicida, which was not previously recorded in Red Sea fish, raising concerns that it could infect native fish.[25]

References

  1. ^ a b c d Pyle, R.; Myers, R. F.; Rocha, L. A.; Craig, M. T. (2010). "Holacanthus ciliaris". IUCN Red List of Threatened Species. 2010: e.T165883A6156566. doi:10.2305/IUCN.UK.2010-4.RLTS.T165883A6156566.en. Retrieved 19 November 2021.
  2. ^ a b c d e Froese, Rainer (December 2019). "Holacanthus ciliaris (Linnaeus, 1758)". Fishbase. Retrieved 21 February 2021.
  3. ^ Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Species in the genus Holacanthus". Catalog of Fishes. California Academy of Sciences. Retrieved 21 February 2021.
  4. ^ "Holacanthus Lacepède, 1802". World Register of Marine Species. Retrieved 24 November 2021.
  5. ^ Scharpf, Christopher; Lazara, Kenneth J. (21 July 2020). "Order Acanthuriformes (Part 1): Families Lobotidae, Pomacanthidae, Drepaneidae and Chaetodontidae". The ETYFish Project Fish Name Etymology Database. Retrieved 22 February 2021.
  6. ^ a b c d e f g h i "Holacanthus ciliaris". Discover Fish. Florida Museum. 10 May 2017. Retrieved 22 February 2021.
  7. ^ a b c Alva-Campbell, Y.; Floeter, S. R.; Robertson, D. R.; Bellwood, D. R.; Bernardi, G. (2010). "Molecular phylogenetics and evolution of Holacanthus angelfishes (Pomacanthidae)". Molecular Phylogenetics and Evolution. 56 (1): 456–461. doi:10.1016/j.ympev.2010.02.014. PMID 20171293.
  8. ^ a b c Tariel, J.; Longo, G. C.; Bernardi, G. (2016). "Tempo and mode of speciation in Holacanthus angelfishes based on RADseq markers". Molecular Phylogenetics and Evolution. 98: 84–88. doi:10.1016/j.ympev.2016.01.010. PMID 26876637. S2CID 205840957.
  9. ^ Reyes-Bonilla, H.; Alvarez-Filip, F.; Sánchez-Alcántara, I. (2010). "New records of the Townsend angelfish (Holacanthus bermudensis X H. ciliaris hybrid) and range extension of the Blue angelfish (H. bermudensis) in the Caribbean Sea". Caribbean Journal of Science. 46 (2–3): 339–345. doi:10.18475/cjos.v46i2.a24. S2CID 130966144.
  10. ^ Feddern, H. (1968). "Hybridization between the western Atlantic angelfishes, Holacanthus isabelita and H. ciliaris" (PDF). Bulletin of Marine Science. 18 (2): 351–382.
  11. ^ a b c Deloach, Ned; Deloach, Anne (2019). Reef Fish Behavior: Florida, Caribbean, Bahamas (2nd ed.). New World Publications. pp. 175–176, 180–181. ISBN 978-1878348685.
  12. ^ Shen, K-N; Chang, C-W; Chen, C-H; Hsiao, C-D (2015). "Complete mitogenomes of King angelfish (Holacanthus passer) and Queen angelfish (Holacanthus ciliaris) (Teleostei: Pomacanthidae)". Mitochondrial DNA Part A. 27 (4): 2815–2816. doi:10.3109/19401736.2015.1053081. PMID 26119118. S2CID 207745744.
  13. ^ "Species: Holacanthus ciliaris, Blue angelfish". Shorefishes of the Greater Caribbean online information. Smithsonian Tropical Research Institute. Retrieved 22 February 2021.
  14. ^ a b c d Snyder, David B.; Burgess, George H. (2016). Marine Fishes of Florida. Johns Hopkins University Press. pp. 221–224. ISBN 978-1421418728.
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Queen angelfish: Brief Summary ( anglais )

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The queen angelfish (Holacanthus ciliaris), also known as the blue angelfish, golden angelfish, or yellow angelfish, is a species of marine angelfish found in the western Atlantic Ocean. It is a benthic (ocean floor) warm-water species that lives in coral reefs. It is recognized by its blue and yellow coloration and a distinctive spot or "crown" on its forehead. This crown distinguishes it from the closely related and similar-looking Bermuda blue angelfish (Holacanthus bermudensis), with which it overlaps in range and can interbreed.

Adult queen angelfish are selective feeders and primarily eat sponges. Their social structure consists of harems which include one male and up to four females. They live within a territory where the females forage separately and are tended to by the male. Breeding in the species occurs near a full moon. The transparent eggs float in the water until they hatch. Juveniles of the species have different coloration than adults and act as cleaner fish.

The queen angelfish is popular in the aquarium trade and has been a particularly common exported species from Brazil. In 2010, the queen angelfish was assessed as least concern by the International Union for Conservation of Nature as the wild population appeared to be stable.

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