If you have the habit of succumbing to "impulse purchases," there is little chance you resisted the temptation to bring home the first dwarf you saw. As one of the most popular fish families amongst home aquarists, the Pomacanthidae, or the marine Angelfish, have become a staple in the hobby as "show" fish. The foremost reason for their popularity is obviously their striking coloration. For the "Fish Tales" of February, I'll select the largest genus from the Pomacanthidae, Centropyge, and discuss some of the more commonly available members of this genus.

The extrmely rare C. boylei is seen here. Enjoy the photo, as they are almost as rare as the specimens themselves. The collection and photography of one is difficult due to the depths of 300 feet or more that are required in order to find one. Photo courtesy of Richard Pyle.

Meet the Family

The Pomacanthidae has been frequently taxonomically revised, and as a whole, needs further investigation. When Fraser-Brunner (1933) elevated Centropyge from sub-genus to generic status, he included multifasciatus as a Centropyge. Even so, some authors (Herre, 1953) still believed Centropyge was a sub-genus of Holacanthus. When Randall and Caldwell described a new Centropyge in 1973, they recognized those same seven genera as did Fraser-Brunner (1933), but concluded multifasciatus "warrants a genus by itself," but stopped short of naming one. One year later Randall revised his previous belief of multifasciatus requiring a monotypic genus, and deduced that multifasciatus belonged in Centropyge. In 1991 Burgess agreed with Randall and Caldwell (1973) and moved multifasciatus into a monotypic genus, Paracentropyge. At the same time Burgess also described a second monotypic genus, and moved venustus from Centropyge to the newly described genus Sumireyakkome. Not long thereafter, Pyle and Randall (1992) disregarded these two monotypic genera and placed venustus and multifasciatus into Centropyge. Fast-forward to present time and Richard Pyle is currently working on his thesis, wherein he is outlining the family Pomacanthidae. Once finished, Pyle will recognize the seven widely agreed upon genera (see below). He will also bring back Paracentropyge and place multifasciatus and boylei into this genus. Venustus is still undetermined, but it is likely to also be found in Paracentropyge as well. He expects many other ichthyologists to disagree with his findings, and thus add further confusion to this family. Coincidently, Burgess does not agree with the seven genera, and he plans on publishing descriptions of new Centropyge in the future. Thus, it is likely the taxonomic state of Centropyge and Pomacanthidae will be turned upside once again. In any case, roughly 75 species are represented within Pomacanthidae.

• Arusetta was originally used for the Red-Sea species asfur. It has not been used in a long time, but there is reason to regard the Pacific species of "Pomacanthus" as distinct from the two Caribbean and one Easter Pacific species of "true" Pomacanthus. If, indeed, the Pacific forms do warrant a distinct genus separate from Pomacanthus, then this will have to be Arusetta (Pomacanthops and Pomacanthodes are not available for this purpose) (Pyle, pers. comm.).
  

Centropyge, commonly called dwarf or pygmy angels, is represented by 33 species (see below) (Note that C. woodheadi is no longer listed as it is now regarded as a synonym of C. heraldi). Like all of their larger cousins, dwarf angels have a continuous dorsal and anal fin. The body is considerably compressed, and has 24 vertebrae. As the family name Pomacanthidae implies (Poma- = operculum, acanth- (Greek) = spine), the operculum has a large, prominent spine. This spine is the easiest way for a hobbyist to determine whether the fish is an angel or not.

For the chart below I will follow Pyle and Randall (1992) and use seven genera of Pomacanthidae. As a result, boylei, multifasciatus, and venustus are listed as Centropyge.

There are also five described natural hybrid Centropyge, and possibly two others awaiting further research and description (see below). All of these natural hybrids have overlapping boundaries, referred to as "hybrid zones." These hybrids are believed to be a result of the rarity of one species, and an abundant number of the other species. Pyle and Randall(1994) believe the rare Centropyge joins the harem as a female, though more conclusive evidence will be required. I will touch on hybrids a little bit more later on. For those of you wishing to understand hybrids to a greater extent, please view the full reference from above, which is linked below in the "References" section.

It is interesting to note that if we follow the Burgess (1991) description you will find the multifasciatus x venustus hybrid actually becomes an inter-generic hybrid between Paracentropyge multifasciatus and Sumireyakko venustus.

There have also been sightings or photographs taken of numerous fish that may represent another natural hybrid, though no specimen was obtainable for further research.

Two other Centropyge species, flavissimus and yrolikii, have overlapping ranges in both the Ogasawara Islands and the Great Barrier Reef/Coral Sea regions, but no hybrid has been discovered (Pyle and Randall, 1994). This bucks the previously set trend of hybrids resulting from shared waters. Possible causes for this are the lack of research done in the areas of overlap, coupled with the lack of collection for home aquaria in these areas.

C. interruptus is a rare import for the hobby, and usually commands a price tag of over $500USD. Photos courtesy of David Wilson.

In the Wild

Dwarf angels are found in both the tropical Atlantic and Pacific Oceans, though the vast majority of them can be found in the tropical western Pacific. They can be found from several feet of water down to several hundred feet deep. In all cases, Centropyge are found amongst coral rubble, branches, and cave systems with quick access to cover. Some species will spend over half of their life hiding within this cover. They are all territorial and haremic (Moyer and Nakazomo, 1978), defending areas as small as 5' x 12' and up to 80 m² (DeLoach, 1999 and Micheal, Coral Realm), which contain from 1 - 6 females (Moyer and Nakazono, 1978 and Moyer, 1990).

The main component of the diet for most dwarf angels is algae and detritus. However, they will consume coral polyps, coral slime, sponges, small invertebrates, and feces from other fish. In some species, up to 50% of their stomach is filled with sand that they have picked up while feeding on detritus (DeLoach, 1999).

In 1978 Moyer and Nakazono showed Centropyge interruptus to be a protogynous hermaphrodite - in other words, they enter the world without a sex. As they mature, they first take on the female sex organs. Similar to other protogynous hermaphrodites, the male is the result of a dominant female that has undergone a sex change due largely to a male that has either died or disappeared in some manner. Shortly thereafter, Randall and Yasuda (1979) believed Centropyge shepardi to be a protogynous hermaphrodite. As research has continued, it is now believed that all Centropyge spp. are protogynous hermaphrodites.

Female dwarf angels establish their den within the center of their territory. Little aggression is exerted towards other fish, the exception being towards other small fish that share the same dietary habits. The male of the harem has a determined route that he continually follows. This route takes him from one female to the next within his harem. As he follows his predetermined route, he stops and visits with each of the females in his harem for a brief period. All dwarf angels mate at sunset, generally within 10 minutes thereafter. As darkness approaches, the male reduces his feeding and his frequency of visits to his females' increases. Eventually, the male will make a complete trip around his territory in less than 5 minutes. Usually, the female waits for the male in nearby sheltered areas. When the female is not present, the male searches for her and, once sighted, they approach each other quickly, with the male flashing his brilliant colors and in some instances, making audible chirping noises. The two fish spend less than five seconds with each, and the male moves onto the next female. With each subsequent visit, the duration of the visit lasts longer. As the moment of truth approaches, the male will swim tight circles around the female before finally moving on. At this point, the female will sometimes follow the male to the next female. It is not rare to have two or three females located at the same site. Finally, when a female is ready, she slowly rises from the sandbed, roughly 12 inches. The male gives the final "ok" by nudging the female with his snout. In less than one second the female will push out over 100 eggs and the male will turn laterally and shower the eggs with sperm. With the male leading the way, they both retreat to cover. The entire process takes between four to six seconds. Pygmy angels are pelagic spawners, and the final tail stroke that sends the fish to cover also sends the (hopefully) fertilized eggs higher into the water column. The spawning process involves only one male and one female per ritual, and once finished the male will move onto his next female. Once he has mated with all of his females, he retires for the evening (DeLoach, 1999).

C. joculator is a hardy angelfish if you don't mind the $400-650 price tag! Photo courtesy of John Boe.

In the Home Aquarium

With a few exceptions, the dwarf angels do well within the confines of home aquaria. Even those that have proven themselves as suitable choices for home aquariums do have a few requirements that a hobbyist must meet before the angels will prosper.

The first concern is a well-aged aquarium with plenty of filamentous algal growth. The growth can be on aquarium rockwork, or the non-viewing sides of the aquarium, or preferably both. Regardless, it should be present if you are to consider your tank ready to house a dwarf angel. The filamentous algae will serve as a large portion of the food source/nutritional value for your newly acquired angel. An aquarium without these algae should be considered unfit for a dwarf angel.

Hopefully, a well-aged aquarium is indicative of a well-aged hobbyist, as well. Dwarf angels are not super-sensitive to water quality, but they are susceptible to disease. If the water parameters sway too far away from ideal, the likelihood of your angel getting sick is above average.

Differing only from C. nox by the small amounts of white and yellow, this C. tibicen is seen shortly after arriving from the wholesaler. Photo by Henry C. Schultz III.

The simple coloration of jet black on this C. nox intrigues some hobbyists. Photo courtesy of Chuck Fiterman.

The obvious next concern would be fellow tank mates. Dwarf angels can get along well with a large assortment of marine fish, though a few should be avoided. Obviously, anything that can swallow it should be avoided. This would include most frogfish and scorpionfish, groupers, and toadfishes. Pygmies can get along fine with more active fish such as large angels, butterflies, and surgeonfish; however, they should be added to the aquarium and settled-in prior to the addition of these faster fish. Generally, dwarf angels will not bother small, more docile fish. The common rule of introducing the most aggressive fish last applies here, so make sure your gobies, jawfish, grammas and dragonettes are in the aquarium first. Many people are curious about mixing species of dwarf angels, or even placing several of the same species in the same aquarium. In general, it is best to leave both of these instances for aquariums over 200 gallons. As an example, the smallest known territory for a given harem is C. argi at 5' x 12' with roughly six or seven specimens per harem. If you were to divide this out, you would find each specimen has a territory close to that of a 55-gallon aquarium. Note that I stated this was the smallest territory known. Some species, such as C. ferrugatus, require a considerable more amount of room. Females of this species will defend a territory of roughly 30m². The males defend a territory even larger at 80m². Note that those numbers are per animal.

When trying to mix dwarf angels, be sure to obtain the smallest animals you can get, and add them at the same time. The goal would be to receive all females, or one male and all the rest females. The smaller the fish, the better the chance of acquiring a female. The dominant female will undergo a sex change and become the male if one is not already present. If two males are added, one will need to be removed before they end up killing each other.

Compatibility chart for members of the genus Centropyge:

Fish	Will Co-Exist	May Co-Exist	Will Not Co-Exist	Notes
Angels, Dwarf	X			When mixed as a single male/several female harem.
Angels, Large	X			Good choice provided the tank is large enough for the angel.
Anthias	X			Good choice in larger tanks.
Assessors		X		Assessors may suffer in smaller aquariums.
Basses			X	Larger Basses may attack smaller angels.
Batfish	X			Good choice.
Blennies	X			Good choice.
Boxfishes	X			Good choice in larger aquariums.
Butterflies	X			Good choice in larger aquariums.
Cardinals	X			Good choice.
Catfish		X		Catfish grow large enough to consume dwarfs.
Comet	X			Good choice in aquariums with plenty of rockwork.
Cowfish	X			Good choice in larger aquariums.
Damsels		X		Damsels and dwarf angels compete for food and territory.
Dottybacks		X		Some dottybacks are best left for species tanks.
Dragonets	X			Will co-exist peacefully.
Drums		X		Small Drums in first.
Eels			X	Not recommended. Most eels will get large enough to consume angels.
Filefish	X			Good choice provided the needs of the Filefish are met.
Frogfish			X	Frogfish will consume angels.
Goatfish	X			Should be good tank mates.
Gobies	X			Good choice. Goby in first.
Grammas	X			Good choice. Grammas in first.
Groupers			X	Groupers will consume dwarf angels.
Hamlets	X			Should be good tank mates.
Hawkfish		X		Some Hawkfishes can become aggressive towards smaller tank mates.
Jawfish	X			Good choice provided the tank has open sandbed for the jawfish and plenty of live rock for the angel.
Lionfish		X		Larger adults can consume dwarf angels.
Parrotfish	X			Parrotfish should ignore dwarf angels, though their adult size might be intimidating to the angel.
Pineapple Fish	X			Should do fine together provided there are plenty of hiding spaces.
Pipefish			X	Pipefish are often delicate and small. Best kept to a species tank.
Puffers		X		Some puffers may be too aggressive for angels.
Rabbitfish	X			Good choice in larger tanks.
Sand Perches		X		Angel in first. Sand Perch may become aggressive once acclimated.
Scorpionfish			X	Adults can consume dwarf angels.
Seahorses			X	Angels may pester the pony, or out-compete it for food.
Snappers			X	Snappers will consume dwarf angels.
Soapfishes			X	Soapfishes may consume smaller dwarf angels.
Soldierfish			X	Adults can consume dwarf angels.
Spinecheeks	X			Should be good tank mates.
Squirrelfish		X		Adults can consume dwarf angels.
Surgeonfish	X			The active swimming of the surgeonfish will keep the dwarf angel close to home.
Sweetlips			X	Can become large enough to consume dwarf angels.
Tilefish		X		The aggressive swimming of Tilefish may keep dwarf angels hiding.
Toadfish			X	Toadfish will consume dwarf angels.
Triggerfish			X	Most Triggerfish are too aggressive for dwarf angels.
Waspfish		X		Angels may pester the Waspfish.
Wrasses	X			Good choice provided the tank is large enough for intended wrasse.

Note: While many of the fish listed are good tank mates for Centropyge, you should research each fish individually before adding it to your aquarium. Some of the fish mentioned are better left in the ocean, or for advanced aquarists.

In regards to mobile invertebrates, you are rather safe placing a dwarf angel in the same tank as them. They will not bother shrimp, cucumbers, larger snails, or crabs. Where everyone seems to be concerned with dwarf angels, however, is in regards to sessile invertebrates. Unfortunately, there is no rule of thumb you can follow or magic potion to add that will guarantee you that your angel will not bother your corals. It is purely hit or miss whether the angel will bother corals. In some instances, the fish will not be nipping at the tissue of the coral, but rather feeding from the coral slime. Regardless, this irritates the coral, and if it is allowed to continue it will likely kill the coral. The best advice I can give you regarding placing dwarf angels in aquariums filled with sessile invertebrates is to be prepared for the worst-case scenario. In some cases the angel will live with you for years and never bother a coral. Sometimes they may nip at the base of corals, but never do any harm. And then sometimes the angel may decide to start nipping at your corals to the point of killing the coral. Regardless what scenario you think you have/will have, BE PREPARED. Dwarf angels are not easily captured. If you remember anything from this column, remember this: if you are not prepared to rip apart your reef at a moment's notice to capture your dwarf angel because it decided it liked the taste of your corals, it is best that you do not consider buying one.

A beautiful C. eibli awaits purchase at a local retailer. Photo by Henry C. Schultz III.

The last considerations you should ponder prior to purchase should be tank size and food. A single dwarf angel can do well in a traditional 55-gallon aquarium, provided the above-mentioned considerations are all met within this tank. The smallest members of the genus, C. argi and C. aurantonotus, will do fine in 40-gallon aquariums. All the dwarf angels will forage from the rockwork, but their diet should be supplemented with prepared foods. Dried algae, also called Nori, are an excellent choice, as are many of the flake foods and frozen foods designed for herbivores. With time, your dwarf angel will accept most any foods offered, including those foods meant for carnivores. So long as your angel receives a wide variety of prepared foods, and has filamentous algae to graze on in the aquarium, their dietary needs will be met.

C. vroliki recieves less attention that its more colorful cousins. Photo by Henry C. Schultz III.

As with all marine fish, only purchase them after a close inspection of the fins, mouth, and tail. Ensure there are no tears or frayed fins, and no red spots or open sores present. Make sure the colors on the fish are bright. Also, make sure the fish is alert and active. Finally, ask to see the fish eat.

The bright orange coloration of this specimen is unfortunately noted to occur in those fish that have been "goosed," what the industry calls a light dosage of cyanide. Specimens like this should be avoided! Photo by Henry C. Schultz III.

Reproduction in the Home Aquarium

Reproduction in the home aquarium has not been entirely successful at this time. There are reports from as far back as the early 1990's of hobbyists having successful spawns. However, raising the fry to adulthood has not been successful by hobbyists. More recently, The Oceanic Institute has had success with both spawning and rearing the fry for C. loriculus. The largest contributing factor was the development of culturing techniques for food that is suitably sized microscopic organisms. This marks a huge achievement for the marine ornamental hobby and hopefully is the start of even larger things to come.

Meet the Species

I'll start the introductions with the smallest members of the family. These little gems are perfect for smaller aquariums where other members of the genus would be a tight fit. All three of the species will not quite reach three inches in length, and they all look fairly similar. They differ mostly by the waters they call home. The Cherub Angelfish, or Centropyge argi, can be found throughout the Caribbean, and into southern Florida. The fish is blue overall, with a yellow face. Centropyge aurantonotus, or the Brazilian Flameback Angelfish, overlaps territory with C. argi near Curacao, as well as extending southward to Brazil. As the common name implies, it has a streak of yellow that starts at the face, similar to C. argi, and runs down the length of the back. Finally, the third look-alike, Centropyge acanthops, hails from Africa. Based on its locale, the common name of African Flameback Angelfish was given. Its coloration is very similar to C. aurantonotus, but differs in that the yellow coloration of C. aurantonotus is orange on C. acanthops.

The smallest of the dwarfs is C. argi. It's small size makes it a good choice for aquariums smaller than 55 gallons. Photo courtesy of Greg Rothschild.

Another of the smallest dwarfs is C. acanthops. Photo courtesy of Greg Rothschild.

One of the most common and popular dwarf angels would be Centropyge bispinosus. Commonly referred to as the Coral Beauty, it is often the first dwarf angel kept by many hobbyists, probably because it is regularly stocked at most aquarium stores and the usual magnetic attraction to dwarf angelfish overwhelms the aquarist. Like all Centropyge spp. it will spend the vast majority of its time within the crevices of the rockwork. It is a typically sized dwarf angel, reaching four inches in length. The regular availability of this fish is probably due to the vast area this species can be found in. Their range extends from Guam to the Philippines, and from Australia to South Africa. The body of the fish is deep purple, and has varying degrees of orange in the center.

Centropyge bispinosus in a home aquarium. Photo courtesy of Sean Tobin.

A dark blue color variant of C. bispinosa (above). A more typically colored specimen is seen below. Photos courtesy of Greg Rothschild above, Sean Tobin below.

The most popular pygmy angel amongst aquarists would have to be the Flame Angel, or Centropyge loriculus. The brightly colored red body of the fish will grab the attention of most any hobbyist. Once again, C. loriculus is an average size for pygmy angels, four inches. It an be found from 2 - 200 feet of water around the Hawaiian Islands, Marshall Islands, and Society Islands. Individuals with a single black stripe behind the head are from the Marquesas Islands. Otherwise, flame angels have three to seven black stripes across the body. All flame angels have a blue trim on the rear of the dorsal and anal fins, with the males having a larger area of blue, as well as more vivid coloration in the blue.

C. loricula in a home aquarium. Photos by Greg Rothschild.

Another Hawaiian native is the Potter's Angel, or Centropyge potteri. It is most common around the Hawaiian Islands, though it can also be found near Johnston Atoll. It has a large depth range, 30 feet to nearly 500 feet, though it isn't found as deep as some of its cousins. The potter's angel is always found in harems ranging from four to eight specimens.

Yet another dwarf angel common around the Hawaiian islands is C. fisheri. Photo courtesy of Henry C. Schultz III.

A trio of yellow dwarf angels is next. When this species hails from Ryukus to the Ducie Islands, the Lemonpeel Angel boasts blue around the eye, gill plate, and gill spike. When from the Indian Ocean around Christmas Island, the blue encircling the eye is absent. It grows slightly larger than the average dwarf, six inches. The Heralds Angelfish, or Centropyge heraldi, it mostly yellow with a small gray patch directly behind the eye. What used to be considered a color morph of C. heraldi has recently been given its own species status - Centropyge woodheadi. The Woodhead's Angelfish differs from C. heraldi only by a black saddle on its dorsal fin. It is distributed around Fiji, Tonga, and the Philippines, while C. heraldi can also be found around southern Japan and the Great Barrier Reef.

The bright yellow of C. heraldi is an attractive option for the smaller aquarium. Photo by Henry C. Schultz III.

Nearly identical to C. heraldi is C. flavissima. The difference should be obvious. Photo by Henry C. Schultz III.

Though not considered rare, the Golden Angelfish is not common in our hobby. This is probably because of its ultra-secretive nature, which behavior generally follows it into home aquariums. Centropyge aurantius can be found in waters shallower than 50 feet across Indonesia, Papua New Guinea, the Solomon Islands, Queensland, Australia and American Samoa. It also exhibits the typical four inch body length of most dwarf angels.

C. aurantius must be seen in person to truly appreciate the remarkable gold colorations. Photo courtesy of Joe Somerville.

One of the largest pygmy angelfish is the Bicolor Angel, or Centropyge bicolor, measuring in at a whopping six inches. It is generally not as hardy as most of the other dwarf angelfish, regularly succumbing to an early exit from aquariums. Harems in the wild include large numbers (for dwarfs) of both mature and immature females. They are most widespread around the Solomon Islands, Fiji Islands, Great Barrier Reef, and the Philippines. This species, as well as any others from this genus, are likely to fare better in aquariums if they are not collected from the Philippine Islands, which still practice unsound collection techniques.

Although not as hardy as some of the other dwarf angels, C. bicolor has a remarkably simplitic beauty to it that garnishes a ton of attraction. Photo courtesy of Greg Rothschild.

The Rusty Angelfish, or Centropyge ferrugatus, is yet another dwarf angel fitting into the four-inch category. It can be found in less than 100 feet of water from southern Japan to Taiwan. Extensive research has gone into the range of this angel, and from this research we know that each female will claim a territory of up to 30m², though this may overlap with other females. The male will stake out over 80m². Unlike other species, rusty angels were noted to form two subgroups of females, each with a dominant female and several smaller or even immature females.

Slightly smaller than the average Centropyge, measuring in at just over 3 inches, is the Multicolor Angelfish, Centropyge multicolor. Although it is common on the reefs of the Marshall, Fiji, and Cook Islands, it usually inhabits water deeper than 100 feet, thereby making collection difficult. Nonetheless, they do show up in the hobby on occasion.

A rare find in the hobby, due largely to collection difficulties, is Centropyge hotumatua. The Easter Island Angelfish is so named because thus far, Easter Island is the only place it has been sighted. Again, it is another smaller pygmy, reaching just over three inches. Usually, the Easter Island pygmy has metallic blue encircling the eye, and the rear dorsal and anal fins are deep blue, almost black.

The two dwarf angelfish within the Pomacanthidae family that have brought about the most grief and discussion among ichthyologists are Centropyge venustus, or the Venustus Angel, and Centropyge multifasciatus, or the Multi-barred Angelfish. Each may reach nearly five inches in length, and are typically found inside of caves and under overhangs. Due to this tendency, these fish are best housed in dimly lit aquariums. Generally, like C. bicolor, these fish have a tough time acclimating to home aquariums. Both can be found in the western Pacific, though C. multifasciatus has a larger home range.

The toxonomic future of C. venustus is still undecided. Photo courtesy of Debbi Edwards.

C. multifaciatus is a member of Centropyge, at least for a little while longer. From the photo it's easy to see how this fish derived its common name: Multi-barred Angelfish. Photo courtesy of Shao Ng.

Conclusion

For those marine aquarium keepers that do not have the luxury of a large aquarium, the pygmy angels enable them the opportunity to maintain individuals of one of the most sought after fish families worldwide. Most of these beautiful fish adapt extremely well in marine aquariums. For the reef keepers, it is a gambling proposition. Will the angel nibble on your prized brain coral, or will it simply ignore all corals? If you play the game and win, the reward will likely be the most attractive fish in your aquarium.

References:

Allen, G.R., 1985 Butterfly and angelfishes of the world, volume 2. Mergus Publishers, Melle, Germany.

Barton, N.H. & K.S. Gale. 1993. Genetic analysis of hybrid zones. pp. 13-45. In: R.G. Harrison (ed.) Hybrid zones and the evolutionary process, Oxford University Press, New York.

Burgess, W.E., 1991 Two new genera of angelfishes, family Pomacanthidae. Tropical Fish Hobbyist , March 1991:68-70.

Herre, A.W.C.T., 1953 Check list of Philippine fishes. Res. Rep. U.S. Fish Wild. Serv., (20):977 p.

Kosaki, R.K. & D. Toyama. 1987. Gold morphs in Centropyge angelfish. Freshwater Mar. Aquar. 10(7): 8-11.

Krupp, F. and H. Debelius, 1990. The hybrid of Centropyge multifasciatus x Holacanthus venustus from the Phillipines and notes on aberrant colour forms of Centropyge multispinis from the Maldives and Red Sea. Revue fr. Aquariol. 17 (2): 52-56.

Lieske, E. and R. Myers, 1994 Collins Pocket Guide. Coral reef fishes. Indo-Pacific & Caribbean including the Red Sea. Haper Collins Publishers, 400 p.

Lutnesky, M.M.F. 1992. Behavioral ecology of reproduction in the pomacanthid angelfish Centropyge potteri. Ph.D. Dissertation, University of Hawaii, Honolulu. 155 pp.

Moyer, J.T. and A. Nakazono, 1978. Population structure, reproductive behavior, and protogynous hermaphroditism in the angelfish Centropyge interruptus at Miyake-jimi, Japan. Japan J. Ich

Moyer, J.T. 1981. Interspecific spawning of the pygmy angelfishes Centropyge shepardi and C. bispinosus at Guam. Micronesica 12(1-2): 119-124.

Pyle, R.L. 1992. Rare and unusual marines: a hybrid angelfish Centropyge flavissimus x eibli. Freshwater Mar. Aquar. 15(3):98-110,212.

Pyle, R.L. 1992. Rare and unusual marines: another hybrid angelfish Centropyge loriculus x potteri. Freshwater Mar. Aquar. 15(8): 40-45.

Pyle, R.L. and J.E. Randall, 1993. A new species of Centropyge from the Cook Islands, with a redescription of Centropyge boylei. Revue fr. Aquariol. 19(4):115-124.

Pyle, R.L and J.E. Randall, 1994. A review of hybridization in marine angelfishes (perciformes: Pomacanthidae). Kluwer Acedemic Publishers. 41:127-145.

Randall, J.E. & F. Yasuda. 1979. Centropyge shepardi, a new angelfish from the Mariana and Ogasawara Islands. Japan. J. Ichthyol. 26: 55-61.

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