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66-491: Holacanthus bermudensis , also known as the blue angelfish or Bermuda blue angelfish , is a species of ray-finned fish, a marine angelfish belonging to the family Pomacanthidae . It occurs in the western Atlantic Ocean. Holacanthus bermudensis was first formally described as Holacanthus ciliaris bermudensis in 1876 by the American ichthyologist George Brown Goode (1851–1896). The original type specimens from Bermuda,

132-413: A cleaning station . It is thought that the vivid, contrasting colours of the juveniles may indicate the establishment of a cleaning station to potential clients. Holacanthus bermudensis isn't important commercially, but it is common in the aquarium trade. Marine angelfish See text Marine angelfish are perciform fish of the family Pomacanthidae . They are found on shallow reefs in

198-399: A combination of both visual and olfactory advertising of the territory. The male ring-tailed lemur has a specialised adaptation to assist in leaving visual/olfactory territorial marks. On their inner forearm (antebrachial) is a scent gland which is covered by a spur . In a behaviour called "spur marking", they grasp the substrate, usually a small sapling, and drag the spur over it, cutting into

264-616: A facultative purpose for this species, in which the more fragrance-rich sites there are, the greater the number of habitable territories. Since these territories are aggregated, females have a large selection of males with whom to potentially mate within the aggregation, giving females the power of mate choice . Similar behaviour is also observed in the Eulaema meriana orchid bee. Males in this species of bee show alternative behaviours of territoriality and transiency. Transient male bees did not defend territories, but instead flew from one territory to

330-525: A good rural habitat, but as large as 300 hectares (740 acres) in a poor habitat. On average, a territory may be approximately 50 hectares (120 acres), with main setts normally at least 500 metres (1,600 ft) apart. In urban areas, territories can be as small as 5 hectares (12 acres), if they can obtain enough food from bird tables, food waste or artificial feeding in suburban gardens. Spotted hyenas ( Crocuta crocuta ) have highly variable territory sizes, ranging from less than 4,000 hectares (9,900 acres) in

396-490: A long-living species and are easy to care for. They were very expensive in the aquarium trade when first discovered, but have become more popular and therefore less pricey. The queen angelfish grows to be 45 cm (18 in). With neon blue and yellow scales and iridescent purple and orange markings, surprisingly it is not conspicuous, and actually hides very well, and is very shy. As juveniles, some species are different colors than when they reach adulthood. For example,

462-412: A nasty neighbour strategy. Group-living male breeders are nearly five times more aggressive towards their neighbours than towards strangers, leading to the prediction that neighbours are the most important competitors for paternity. Using a molecular parentage analysis it has been shown that 28% of offspring are sired by neighbouring males and only 7% by strangers. In certain species of butterflies, such as

528-428: A perspective. Several other type of resource may be defended including partners, potential mates, offspring, nests or lairs, display areas or leks . Territoriality emerges where there is a focused resource that provides enough for the individual or group, within a boundary that is small enough to be defended without the expenditure of excessive effort. Territoriality is often most strong towards conspecifics , as shown in

594-588: A single male dominant over several females. As juveniles, some species may eke out a living as cleaner fish . Common to many species is a dramatic shift in coloration associated with maturity. For example, young male ornate angelfish , Genicanthus bellus , have broad, black bands and are indistinguishable from females; as they mature, bright orange bands develop on the flanks and back. Thought to correspond to social rank, these colour shifts are not necessarily confined to males; all marine angelfish species are known to be protogynous hermaphrodites . This means that if

660-588: A strategy termed the dear enemy effect in which two neighbouring territorial animals become less aggressive toward one another once territorial borders are well-established and they are familiar to each other, but aggression toward unfamiliar animals remains unaffected. The converse of this is the nasty neighbour effect in which a territory-holder shows heightened aggression toward neighbouring territory-holders but unaffected aggression to unfamiliar animals or distant territory-holders. These contrasting strategies depend on which intruder (familiar or unfamiliar) poses

726-506: A vivid yellow caudal , pectoral and pelvic fins . They also have a number of vertical white bars on the body with bright blue margins to the dorsal and anal fins . The adults are bluish yellowish on the body with a vivid yellow face. They have blue highlights on the chest and forehead and blue and yellow pectoral fins while the caudal fin has yellow margins. The dorsal and anal fins also have yellow margins and long yellow streamers. The dorsal fin contains 15 spines and 19–21 soft rays while

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792-450: Is a lekking reptile. Males start to establish small display territories two months ahead of the mating season. Rather than retaining a territory simply by fighting, for some animals this can be a 3-stage process. Many animals create "sign-posts" to advertise their territory. Sometimes these sign-posts are on the boundary thereby demarcating the territory, or, may be scattered throughout the territory. These communicate to other animals that

858-421: Is a behaviour used by animals to identify their territory. Most commonly, this is accomplished by depositing strong-smelling substances contained in the urine , faeces , or, from specialised scent glands located on various areas of the body. Often, the scent contains pheromones or carrier proteins such as the major urinary proteins to stabilize the odours and maintain them for longer. The animal sniffing

924-520: Is a benthic species, living close to the seabed where there are areas of sponges , coral, or rock at depths between 2 and 93 metres (6.6 and 305.1 ft). It is a diurnal species, active in daylight and hiding within the reef at night. Juveniles prefer more sheltered area such as bays, channels, and inshore reefs. This species has a diet that comprises largely of sponges although they have been occasionally recorded feeding on tunicates , corals, and algae . The juveniles act as cleaner fish , feeding on

990-523: Is a large (up to 8 cm in length) limpet. It lives in association with an approximately 1,000 cm^2 area of algal film in which its grazing marks can be seen, whereas the remainder of the rock surface is usually free of any visible film. These areas of algal film represent the territories of the Lottia; within them the animals do all their grazing. They keep their territories free of other organisms by shoving off any intruders: other Lottia, grazing limpets of

1056-484: Is a series of stylised postures, vocalisations, displays, etc. which function to solve the territory dispute without actual fighting as this could injure either or both animals. Ritualized aggression often ends by one of the animals fleeing (generally the intruder). If this does not happen, the territory may be defended by actual fighting, although this is generally a last resort. Scent marking, also known as territorial marking or spraying when this involves urination ,

1122-406: Is attached to a large yolk sac but which has no functioning fins, eyes or gut. After around 48 hours the yolk is absorbed, while it is absorbed the pro-large changes into a true larvae and commences eating plankton. Subsequent growth is quick and in the 3 to 4 weeks following their hatching they settle on the seabed. Juveniles are highly territorial ,they defend territories where they have established

1188-636: Is different from normal urination, which is done while squatting. This posture is exclusive to alpha wolves of either sex, although the alpha male does this most often. The alpha female usually urinates on a scent post that her breeding partner has just urinated on, although during the mating season, the female may first urinate on the ground. All other females in the pack, and also young wolves and low-ranking male wolves, urinate while squatting. Similar urination postures are used by coyotes and golden jackals . Males and female ring-tailed lemurs ( Lemur catta ) scent-mark both vertical and horizontal surfaces at

1254-457: Is laid in well defined piles. There may be 20 to 30 of these piles to alert passing rhinoceroses that it is occupied territory. Other males may deposit dung over the piles of another and subsequently the sign-post grows larger and larger. Such a dung heap can become up to five metres wide and one metre high. After defecating, greater one-horned rhinos scratch their hind feet in the dung. By continuing to walk, they "transport" their own smell around

1320-554: Is least likely with insectivorous birds, where the food supply is plentiful but unpredictably distributed. Swifts rarely defend an area larger than the nest. Conversely, other insectivorous birds that occupy more constrained territories, such as the ground-nesting blacksmith lapwing may be very territorial, especially in the breeding season during which they not only threaten or attack many kinds of intruders, but have stereotyped display behaviour to deter conspecifics sharing neighbouring nesting spots. The owl limpet ( Lottia gigantea )

1386-492: Is not always a fixed behavioural characteristic of a species. For example, red foxes ( Vulpes vulpes ) either establish stable home ranges within particular areas or are itinerant with no fixed abode . Territories may vary with time (season). For example, European robins defend territories as pairs during the breeding season but as individuals during the winter. Resource availability may cause changes in territoriality. For example, some nectarivores defend territories only during

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1452-399: Is reversed. Animals may use several strategies to defend their territories. The first game theory model of fighting is known as the hawk-dove game . This model pits a hawk strategy (always try to injure your opponent and only withdraw from the contest if an injury is received) against a dove strategy (always use a non-injurious display if the rival is another dove and always withdraw if

1518-477: The Australian painted lady butterfly and the speckled wood butterfly , the male defends territories that receptive females are likely to fly through such as sunny hilltops and sunspots on a forest's floor. Territory defence in male variegated pupfish ( Cyprinodon variegatus ) is dependent on the presence of females. Reduced aggression consistent with the dear enemy effect occurs between conspecific neighbours in

1584-808: The Ngorongoro Crater to over 100,000 hectares (250,000 acres) in the Kalahari . In birds, golden eagles ( Aquila chrysaetos ) have territories of 9,000 hectares (22,000 acres), least flycatchers ' ( Empidonax minimus ) territories are about 600 square metres (6,500 sq ft) and gulls have territories of only a few square centimetres in the immediate vicinity of the nest. Territories can be linear. Sanderlings ( Calidris alba ) forage on beaches and sandflats. When on beaches, they feed either in flocks or individual territories of 10 to 120 metres of shoreline. The time to develop territories varies between animals. The marine iguana ( Amblyrhynchus cristatus )

1650-552: The Uganda kob (a grazing antelope ) and the marine iguana, males defend the lek site which is used only for mating. Many species demonstrate polyterritoriality, referring to the act of claiming or defending more than one territory. In the European pied flycatcher ( Ficedula hypoleuca ), researchers assert that males exhibit polyterritoriality to deceive females of the species into entering into polygynous relationships. This hypothesis, named

1716-472: The butterflyfishes , a related family of similarly showy reef fish. Marine angelfish are distinguished from butterflyfish by the presence of strong preopercle spines (part of the gill covers) in the former. This feature also explains the family name Pomacanthidae; from the Greek πομα, poma meaning "cover" and ακάνθα, akantha meaning "thorn". Many species of marine angelfishes have streamer-like extensions of

1782-458: The ectoparasites picked from the skin of other fishes visiting communal cleaning stations. They have the ability to produce loud thumping sounds which are thought to startle predators and draw the attention of conspecifics. The adults of the blue angelfish are usually encountered in pairs and they remain in these pairs all year. This has been interpreted as meaning that they are monogamous . When breeding they spawn by slowly swimming upwards in

1848-599: The syntypes , were set aside and replaced by a neotype , which was the holotype of Angelichthys isabelita which had been described by David Starr Jordan and Cloudsley Louis Ritter with the type locality given as Key West in Florida. This is set out in the Official List of Specific Names in Zoology ( ICZN , Opinion 2003). This species frequently hybridises with the sympatric queen angelfish ( Holacanthus ciliaris) . H. isabelita

1914-854: The Blue Angelfish is a vibrant, electric blue color with black and white stripes or spots. When they reach adulthood, they turn a grayish color with yellow and blue fins and dark spots on their bodies. The larger species are also quite bold and seemingly fearless; they are known to approach divers. While the majority adapts easily to captive life, some are specialist feeders which are difficult to maintain. Feeding habits can be strictly defined through genus, with Genicanthus species feeding on zooplankton and Centropyge preferring filamentous algae . Other species focus on sessile benthic invertebrates ; sponges , tunicates , bryozoans , and hydroids are staples. On Caribbean coral reefs, angelfishes primarily eat sponges, and have an important role in preventing

1980-490: The Centropyge genus, are range restricted or endemic to specific islands or small island groups. Territorial (animal) Territoriality is only shown by a minority of species. More commonly, an individual or a group of animals occupies an area that it habitually uses but does not necessarily defend; this is called its home range . The home ranges of different groups of animals often overlap, and in these overlap areas

2046-408: The ability of an animal to survive and raise young. The proximate functions of territory defense vary. For some animals, the reason for such protective behaviour is to acquire and protect food sources, nesting sites, mating areas, or to attract a mate. Among birds, territories have been classified as six types. Reports of territory size can be confused by a lack of distinction between home range and

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2112-501: The absence of females, but the presence of a female in a male's territory instigates comparably greater aggression between the neighbours. In the Skylark ( Alauda arvensis ), playbacks of neighbour and stranger songs at three periods of the breeding season show that neighbours are dear enemies in the middle of the season, when territories are stable, but not at the beginning of the breeding season, during settlement and pair formation, nor at

2178-554: The anal fin has 3 spines and 20–21 soft rays. This species attains a maximum total length of 45 centimetres (18 in). Holacanthus bermudensis is found in the western Atlantic from North Carolina to Bermuda , into the Bahamas and Florida to the Gulf of Mexico , and also to Yucatan , Mexico in the west and east to Puerto Rico and the Virgin Islands . Holacanthus bermudensis

2244-432: The animal is present, but can travel long distances and over varied habitats. Examples of animals which use auditory signals include birds, frogs and canids. Wolves advertise their territories to other packs through a combination of scent marking and howling. Under certain conditions, wolf howls can be heard over areas of up to 130 km (50 sq mi). When howling together, wolves harmonize rather than chorus on

2310-582: The bird (i.e. badge) is highly visible when it sings (vocal marking) at the boundary of its territory. The ring-tailed lemur ( Lemur catta ) advertises its territory with urine scent marks. When it is urinating for marking purposes, it holds its extremely distinctive tail high in the air adding a visual component to the advertisement; when it is urinating for eliminative purposes, its tail is only slightly raised. Rhinoceros have poor vision but may use visual marking. Dominant white rhino bulls mark their territory with faeces and urine (olfactory marking). The dung

2376-434: The body with urine), to communicate. Many ungulates , for example the blue wildebeest , use scent marking from two glands, the preorbital gland and a scent gland in the hoof . Territorial scent marking may involve behaviours specific to this activity. When a wolf marks its territory , it lifts a hind leg and urinates on a scent post (usually an elevated position like a tree, rock, or bush). This raised leg urination

2442-488: The case of redlip blenny . This is because the conspecifics share exactly the same set of resources. Several types of resource in a territory may be defended. Food: Large solitary (or paired) carnivores, such as bears and the bigger raptors require an extensive protected area to guarantee their food supply. This territoriality only breaks down when there is a glut of food, for example when grizzly bears are attracted to migrating salmon . Food related territoriality

2508-448: The defended territory. The size and shape of a territory can vary according to its purpose, season, the amount and quality of resources it contains, or the geography. The size is usually a compromise of resource needs, defense costs, predation pressure and reproductive needs. Some species of squirrels may claim as much as 10 hectares (25 acres) of territory. For European badgers , a home range may be as small as 30 hectares (74 acres) in

2574-427: The dominant male of a harem is removed, a female will turn into a functional male. As pelagic spawners, marine angelfishes release many tiny buoyant eggs into the water which then become part of the plankton . The eggs float freely with the currents until hatching, with a high number falling victim to planktonic feeders. In aquariums, two fish usually will breed within their community but will harass other fish in

2640-571: The end, when bird density increases due to the presence of young birds becoming independent. Thus, this dear enemy territoriality relationship is not a fixed pattern but a flexible one likely to evolve with social and ecological circumstances. Some species of bees also exhibit territoriality to defend mating sites. For example, in Euglossa imperialis , a non-social bee species, males have been observed to occasionally form aggregations of fragrance-rich territories, considered to be leks. These leks serve only

2706-430: The genus Acmaea, predatory snails, and sessile organisms such as anemones and barnacles. Nests and offspring: Many birds, particularly seabirds, nest in dense communities but are nonetheless territorial in defending their nesting site to within the distance they can reach while brooding. This is necessary to prevent attacks on their own chicks or nesting material from neighbours. Commonly the resulting superimposition of

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2772-489: The greatest threat to the resident territory-holder. In territory defence by groups of animals, reciprocal altruism can operate whereby the cost to the benefactor in helping defend the territory is less than the gains to the beneficiary. An animal chooses its territory by deciding what part of its home range it will defend. In selecting a territory, the size and quality play crucial roles in determining an animal's habitat. Territory size generally tends to be no larger than

2838-579: The ground nearby, thereby leaving a visual advertisement of the territory. This includes domestic dogs . Several species scratch or chew trees leaving a visual mark of their territory. This is sometimes combined with rubbing on the tree which may leave tufts of fur. These include the Canada lynx ( Lynx canadensis ) and the American black bear ( Ursus americanus ). Many animals have scent glands in their paws or deposit fur during tree-marking, so tree-marking may be

2904-442: The groups tend to avoid each other rather than seeking to confront and expel each other. Within the home range there may be a core area that no other individual group uses, but, again, this is as a result of avoidance. The ultimate function of animals inhabiting and defending a territory is to increase the individual fitness or inclusive fitness of the animals expressing the behaviour. Fitness in this biological sense relates to

2970-426: The mornings when plants are richest in nectar. In species that do not form pair bonds, male and female territories are often independent, i.e. males defend territories only against other males and females only against other females. In this case, if the species is polygynous , one male territory probably contains several female territories, while in some polyandrous species such as the northern jacana , this situation

3036-566: The most combative spiders have the largest territories. Some species of penguin defend their nests from intruders trying to steal the pebbles from which the nest is constructed. Mating opportunities: The striped mouse ( Rhabdomys pumilio ) is group living with one single breeding male and up to 4 communally breeding females per group. Groups typically contain several philopatric adult sons (and daughters) that are believed not to breed in their natal group and all group members participate in territorial defence. Males defend their territory using

3102-418: The nest site entrance to establish their territory. Wombats use feces to mark their territory. They have evolved specialized intestinal anatomy to produce cubical feces to ensure the feces do not roll away. Visual sign-posts may be a short-term or long-term mode of advertising a territory. Short-term communication includes the colouration or behaviour of the animal, which can only be communicated when

3168-407: The organism requires to survive, because defending a larger territory incurs greater energy, time and risk of injury costs. For some animals, the territory size is not the most important aspect of territoriality, but rather the quality of the defended territory. Behavioural ecologists have argued that food distribution determines whether a species is territorial or not, however, this may be too narrow

3234-715: The other. They also did not engage in physical contact with the territorial males. On the other hand, territorial males patrolled an area around a tree and used the same territory for up to 49 days. It also appeared that they gave up territories to new males without violence. Males defend territories solely for mating, and no other resources such as fragrances, nests, nest construction materials, nectar, or pollen are found at these territories. Although most territories contain multiple (potential) resources, some territories are defended for only one purpose. European blackbirds may defend feeding territories that are distant from their nest sites, and in some species that form leks, for example in

3300-498: The overgrowth of reef-building corals by eating faster-growing sponge species. Most marine angelfishes restrict themselves to the shallows of the reef, seldom venturing deeper than 50 m (160 ft). The recently described Centropyge abei is known to inhabit depths of 150 m (490 ft). They are diurnal animals, hiding amongst the nooks and crevices of the reef by night. Some species are solitary in nature and form highly territorial mated pairs; others form harems with

3366-476: The overlaps in their home ranges using their anogenital scent glands . To do this, they perform a handstand to mark vertical surfaces, grasping the highest point with their feet while applying the scent. In the Eastern carpenter bee, Xylocopa virginica , both sexes have glands that evolved for marking the nest. Males, although they have the gland, are unable to produce the marking substance. Females secrete it near

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3432-407: The paths, thus establishing a scent-marked trail. Another method of visually marking their territory is wiping their horns on bushes or the ground and scraping with the feet, although this is likely combined with the smell of the marking animal. The territorial male scrape-marks every 30 m (98 ft) or so around its territory boundary. After leaving a urination mark, some animals scrape or dig

3498-724: The resident is present. Other animals may use more long-term visual signals such as faecal deposits, or marks on the vegetation or ground. Visual marking of territory is often combined with other modes of animal communication. Some animals have prominent "badges" or visual displays to advertise their territory, often in combination with scent marking or auditory signals. Male European robins are noted for their highly aggressive territorial behaviour. They attack other males that stray into their territories, and have been observed attacking other small birds without apparent provocation. Such attacks sometimes lead to fatalities, accounting for up to 10% of adult robin deaths in some areas. The red breast of

3564-448: The rest of the family. The smaller species are popular amongst aquarists , whereas the largest species are occasionally sought as a food fish; however, ciguatera poisoning has been reported as a result of eating marine angelfish. Angelfish vary in color and are very hardy fish. When kept in aquariums they can easily adapt to pH and hardness changes in water and can handle conditions that are not considered to be perfect. They are usually

3630-418: The risk of injury. This is ritualized aggression . Such defense frequently involves a graded series of behaviours or displays that include threatening gestures (such as vocalizations, spreading of wings or gill covers, lifting and presentation of claws, head bobbing, tail and body beating) and finally, direct attack. Territories may be held by an individual, a mated or unmated pair, or a group. Territoriality

3696-408: The rival is a hawk). Another strategy used in territory defence is the war of attrition . In this model of aggression, two contestants compete for a resource by persisting while constantly accumulating costs over the time that the contest lasts. Strategically, the game is an auction in which the prize goes to the player with the highest bid, and each player pays the loser's low bid. Some animals use

3762-514: The same note, thus creating the illusion of there being more wolves than there actually are. Wolves from different geographic locations may howl in different fashions: the howls of European wolves are much more protracted and melodious than those of North American wolves, whose howls are louder and have a stronger emphasis on the first syllable. Animals use a range of behaviours to intimidate intruders and defend their territories, but without engaging in fights which are expensive in terms of energy and

3828-460: The scent frequently displays a flehmen response to assist in detecting the mark. Scent marking is often performed by scent rubbing in many mammals. In many mammal species, scent marking is more frequent during the breeding season . Bears and felids such as leopards and jaguars scent-mark by urinating on or rubbing against vegetation. Prosimians and New World monkeys also use scent marking, including urine washing ( self-anointing

3894-511: The short-range repulsion onto the long-range attraction characteristically leads to the well-known roughly hexagonal spacing of nests. One gets a similar hexagonal spacing resulting from the territorial behaviour of gardening limpets such as species of Scutellastra . They vigorously defend their gardens of particular species of algae, that extend for perhaps 1–2 cm around the periphery of their shells. The desert grass spider, Agelenopsis aperta , often engages in fights over its territory and

3960-407: The soft dorsal and anal fins . The fish have small mouths, relatively large pectoral fins , and rounded to lunate tail fins. The largest species, the gray angelfish , Pomacanthus arcuatus , may reach a length of 60 cm (24 in); at the other extreme, members of the genus Centropyge do not exceed 15 cm (5.9 in). A length of 20 to 30 cm (7.9 to 11.8 in) is typical for

4026-565: The tank, so it is best they have their own with plenty of room. The Pomacanthidae is frequently placed within the large order Perciformes but taxonomists have also placed the family within the order Acanthuriformes, alongside the Chaetodontidae and Acanthuridae, among others. Other authorities have resolved the family as incertae sedis . There are 88 species in eight genera: The more speciose genera are, generally speaking, widely distributed, however some species, especially of

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4092-449: The territory is occupied and may also communicate additional information such as the sex, reproductive status or dominance status of the territory-holder. Sign-posts may communicate information by olfactory , auditory, or visual means, or a combination of these. If an intruder progresses further into the territory beyond the sign-posts and encounters the territory-holder, both animals may begin ritualized aggression toward each other. This

4158-456: The tropical Atlantic , Indian , and mostly western Pacific Oceans . The family contains seven genera and about 86 species. They should not be confused with the freshwater angelfish , tropical cichlids of the Amazon Basin . With their bright colours and deep, laterally compressed bodies, marine angelfishes are some of the more conspicuous residents of the reef . They most closely resemble

4224-401: The water column, moving their abdomens together, and releasing copious amounts of ova and milt . The female may lay between 25 and 75 thousand eggs at a time and up to 10 million eggs in each spawning season. Each of the transparent , pelagic eggs contains a small quantity of oil as a buoyancy aid. The eggs take 15 to 20 hours to hatch, the hatchlings being a type of pre-larval stage which

4290-474: The wood and spreading the gland's secretions. When on the ground, ring-tailed lemurs preferentially mark small saplings and when high in the trees, they usually mark small vertical branches. European wildcats ( Felis silvestris ) deposit their faecal marks on plants with high visual conspicuousness that enhances the visual effectiveness of the signal. Many animals use vocalisations to advertise their territory. These are short-term signals transmitted only when

4356-409: Was formerly considered to be a separate species but is now regarded as a synonym of H. bermudensis . Holacanthus bermudensis has a deep, oval-shaped body that is strongly lateral compressed. It has a short snout with a small mouth that contains small teeth, like the bristles of a brush. The juveniles are yellowish on the anterior part of the body changing to brownish-yellow halfway along. They have

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