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
extremely rare C. boylei is seen here. Enjoy
the photo, as they are almost as rare as the specimens
themselves. Their collection and photography 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.
Genera of Pomacanthidae:
Soon to be Accepted
Genera of Pomacanthidae (Pyle, pers. comm.):
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
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
that are Probable Hybrids:
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.
Hybrids for Centropyge:
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.
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).
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.
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
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.
chart for members of the genus Centropyge:
Will Not Co-Exist
When mixed as a single male/several female harem.
Good choice provided the tank is large enough for the
Good choice in larger tanks.
Assessors may suffer in smaller aquariums.
Larger Basses may attack smaller angels.
Good choice in larger aquariums.
Good choice in larger aquariums.
Catfish grow large enough to consume dwarfs.
Good choice in aquariums with plenty of rockwork.
Good choice in larger aquariums.
Damsels and dwarf angels compete for food and territory.
Some dottybacks are best left for species tanks.
Will co-exist peacefully.
Small Drums in first.
Not recommended. Most eels will get large enough to
Good choice provided the needs of the Filefish are
Frogfish will consume angels.
Should be good tank mates.
Good choice. Goby in first.
Good choice. Grammas in first.
Groupers will consume dwarf angels.
Should be good tank mates.
Some Hawkfishes can become aggressive towards smaller
Good choice provided the tank has open sandbed for
the jawfish and plenty of live rock for the angel.
Larger adults can consume dwarf angels.
Parrotfish should ignore dwarf angels, though their
adult size might be intimidating to the angel.
Should do fine together provided there are plenty of
Pipefish are often delicate and small. Best kept to
a species tank.
Some puffers may be too aggressive for angels.
Good choice in larger tanks.
Angel in first. Sand Perch may become aggressive once
Adults can consume dwarf angels.
Angels may pester the pony, or out-compete it for food.
Snappers will consume dwarf angels.
Soapfishes may consume smaller dwarf angels.
Adults can consume dwarf angels.
Should be good tank mates.
Adults can consume dwarf angels.
The active swimming of the surgeonfish will keep the
dwarf angel close to home.
Can become large enough to consume dwarf angels.
The aggressive swimming of Tilefish may keep dwarf
Toadfish will consume dwarf angels.
Most Triggerfish are too aggressive for dwarf angels.
Angels may pester the Waspfish.
Good choice provided the tank is large enough for intended
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.
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
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.
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
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.
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.
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.
bispinosus in a home aquarium. Photo courtesy of
dark blue color variant of C. bispinosa (above).
A more typically colored specimen is seen below.
Photos courtesy of Greg Rothschild above, Sean Tobin
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.
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.
will only get in the way when describing the beauty
of C. potteri. Photos by Greg Rothschild.
another dwarf angel common around the Hawaiian islands
is C. fisheri. Photo courtesy of Henry C. Schultz
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
bright yellow of C. heraldi is an attractive
option for the smaller aquarium. Photo by Henry C. Schultz
identical to C. heraldi is C. flavissima.
The difference should be obvious. Photo by Henry C.
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.
aurantius must be seen in person to truly appreciate
the remarkable gold colorations. Photo courtesy of Joe
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.
not as hardy as some of the other dwarf angels, C.
bicolor has a remarkably simplistic beauty to it
that garnishes a ton of attraction. Photo courtesy of
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 three 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 its collection
difficult. Nonetheless, they do show up in the hobby on occasion.
multicolor is sometimes confised with C. venusutus.
Photos by Greg Rothschild.
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.
toxonomic future of C. venustus is still undecided.
Photo courtesy of Debbi Edwards.
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.
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.
you have any questions about this article, please visit my author
forum on Reef Central.
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