Cichlids of the New World Part IV - Geophagines
by Dean Hougen
As published in Aqua News September/October 1994
A Publication of the Minnesota Aquarium Society
Aquarticles
Among the North American cichlids, which I covered in Part Three of this series, there
are cichlids with specialized adaptations which allow them to sift food items from the
substrate. As I said there, the members of the Thorichthys section of
'Cichlasoma', and some of the members of the Amphilophus section, have elongated
snouts which point downward and have the mouth positioned terminally, and have eyes which
are set back on the head away from the mouth. These adaptations allow these fish to push
the mouth down deeply into the substrate without endangering their fragile eyes.
In South America there are cichlids which have evolved even further specialization for
sifting for food. These cichlids are the Geophagines, which literally means the
"Earth-eaters," and these fish are the subject of this part of the series.
Of course, Geophagines do not literally eat the earth, they merely sift through it.
This latter, however, they do with great efficiency. Besides adaptations to the facial
structure to allow for rooting about in the substrate, Geophagines have a unique gill
morphology that allows them to easily separate inedible sand from food. From a single
mouthful of mixed sand and food, the edible portion is ingested while the grit is spit
back out of the mouth or passed out behind the gill covers.
Geophagines are also interesting subjects due to their reproductive strategies. There
are a wide variety of reproductive strategies found within the group as a whole, including
biparental substrate-spawning and delayed mouthbrooding (both of which are found in the
Cichlasomines, as discussed in the first three parts of the series); polygynous, immediate
mouthbrooding; harem polygynous, substrate-spawning; and apparently even the
extraordinarily rare, monogamous, immediate mouthbrooding.
Further, whereas reproductive strategies are generally uniform within a given genus, a
range of strategies are present within some individual Geophagine genera.
The Genera
Geophagus Heckel 1840
Like several genera discussed in the earlier parts of this series, the genus Geophagus
has recently been restricted to exclude many of the species that once went by that name.
The species that remain may be thought of by aquarists as the surinamensoids, as the few
members of the restricted Geophagus that are regularly found in the hobby are
almost uniformly referred to as Geophagus surinamensis. According to the current
understanding of these species, however, it is unlikely that the true G. surinamensis has
ever been found in the hobby or that it will be commercially collected in the near future,
due to the location and small extent of its natural range. The species referred to in the
hobby as G. surinamensis are, in reality, other species of the restricted Geophagus,
most commonly G. brachybranchus, G. megasema, and G. proximus.(1)
These fish can easily be distinguished by markings on the face, flank, and caudal fins.
The Geophagus species commonly found in the hobby grow quite large (up to a
foot, total length), but some of the less well-known, and recently described species
apparently stay quite a bit smaller (e.g. up to 6" for G. taeniopareius, and
less than 4.5" for G. grammepareius). Geophagus is known to include
substrate spawners (such as the recently described G. argyrostictus), delayed
mouthbrooders (e.g. G. brachybranchus), and advanced mouthbrooders (e.g. G.
megasema). There are probably at least fifteen species of Geophagus, although
less than a dozen have been described.
Satanoperca Gunther 1862
One of the best known sifters in the aquarium hobby is the fish usually referred to as
"Geophagus jurupari". According to current understanding, however, this
fish and its close relatives really belong in the genus Satanoperca. Further, the
hobby "Jurupari," which possesses white spots on its face and gill covers, is
really S. leucosticta (the species name referring to the white spots). The true S.
jurupari lacks any sort of spotting on its head. These species are delayed
mouth-brooders.
Closely related to these two species are S. pappaterra, and apparently several
other species which either need to be described or need to be matched with names which
have previously been treated as junior synonyms for S. jurupari. These include a
species which resembles S. pappaterra and hails from Peru (2),
a species which resembles S. leucosticto but lacks spots on the gill covers (3),
a species found in Peru which resembles S. jurupari but has a shorter snout (4),
and a comparatively small Satanoperca species which apparently is an immediate
mouthbrooder.(5) Due to its origin, it has been speculated
that this last fish may match with S. mapiritensis.(6)
Satanoperca also contains a complex of sifters with large spots on their sides
and trailers on the last several rays of their dorsal fins. The best known of these
species is S. daemon, which possesses two large spots on its side and a
well-ocellated spot on its caudal peduncle. S. daemon is a substrate spawner with
a twist. It buries its eggs under two inches of sand, digs them up when they hatch only to
bury the wrigglers again, and finally digs up the fry when they are ready to be
free-swimming.
The other spotted Satanoperca species (S. lilith, which has a single
side spot located high up on the side but otherwise resembles S. daemon, and S.
acuticeps, which has three side spots and no ocellus on the caudal spot) have not
been spawned in captivity.
Satanoperca species tend to grow slowly, but most can get quite large -- up to
a foot in length.
Hump-head Eartheaters
The fish I am referring to here as hump-head eartheaters, like the species of Satanoperca
above, were regarded as belonging to Geophagus until that genus was restricted
recently. Unlike the Satanoperca species, however, this group of species is of
"orphaned," that is, temporarily lacking a proper generic placement.
The best known species of this group in the aquarium hobby is 'Geophagus'
steindachneri, the "Red-Hump Eartheater." This fish has gone by many names
in the aquarium hobby, including G. magdalenae, G. hondae (both invalid junior
synonyms for G. steindachneri) and G. pellegrini. 'G.' pellegrini,
properly refers to a different species in the bump-head group, and the other described
species in this complex is 'G.' crassilabris.
These fish come from the northern flowing rivers of Columbia and from Panama. All are
immediate mouthbrooders, and the males like so many African rift-lake cichlids, have no
parental role to play once fertilization has taken place.
Brasiliensoids
Another group of orphaned species formerly belonging to Geophagus are the
brasiliensoids. Currently there is only one recognised species in this group, 'Geophagus'
brasiliensis, which is ostensibly a common aquarium fish. However, once a modern
review of these fish takes place I believe that a number of separate species will be
recognized within this group.
Brasiliensoids known in the hobby are biparental, substrate spawners and are notably
more aggressive than most other Geophagines.
Wayne Leibel has speculated that the brasiliensoids are closely related to the rhabdotus
complex species of Gymnogeophagus, but I think that the unique skeletal
characteristics of that genus (see below) preclude a merger of these species and those at
the generic level.
Gymnogeophagus Ribeiro 1918
The genus name Gymnogeophagus means, literally, naked earth-eater. The prefix
gymno (naked) refers to the fact that these fish lack scales on their cheeks and was the
primary character used to establish the genus in 1918. Since then many ichthyologists have
considered this character as too weak to justify a genus and counted these species as
belonging to Geophagus. In 1976, however, Gosse re-examined these fish and
discovered that they possessed unique skeletal structures at the front of the dorsal fin
including an 'antrorse spine' that is not found in any other new world cichlid. While the
genus Gymnogeophagus contains distinct groups within it and may at some point be
split into distinct genera or subgenera, these skeletal features ensure that these species
will always be set apart from all other cichlids.
The type species of Gymnogeophagus is Gg. balzanii, an unmistakable
cichlid with a steep cranial profile, especially in mature males in which the face may
rise straight up from the mouth to the top of the head, and nearly unique vertical stripes
on the flanks which form double "pin stripes" on the lower side. (This latter
characteristic is shared with a Biotodoma species. See below.) Gg. balzanii is
a harem polygynous, delayed maternal mouthbrooder. That is to say, a single male holds a
breeding territory in which several females care for the eggs and fry and the females
carry the fry in their mouths, though not the eggs.
Quite unlike Gg. balzanii are a group of species regarded as belonging to
Gymnogeophagus, but resembling 'G.' brasiliensis more in overall shape and
coloration. The best known species in this complex is Gg. rhabdotus, although
many fish sold under this name are not this fish at all. As I have shown elsewhere (7),
many of the fish sold locally under this name are really brasiliensoids as they lack the
unique skeletal features of Gymnogeophagus. Paul Loiselle has written widely
about Gg. rhabdotus under the name Gg. australis (8),
but it is now known that that species is a member of the gpnnogenys complex (9)
(see below).
Gg. rhabdotus is known to be a simple substratum spawner. The other two
members of the rhabdotus complex are Gg. meridionalis and Gg.
setequedas.
Finally, the remainder of the fish in Gymnogeophagus belong to the gwnnogenys
complex. Like Gg. rhabdotus, many fish sold as Gg. gymnogenys really
were (are) not. Finally, however, the true Gg. gymnogenys appears to be getting a
hold in the hobby, so it is possible to get the real thing these days. Gg. gymnogenys
is an elongated sifter and adult males are unmistakable. They posses red unpaired fins, a
red patch on the gill covers, and unique black markings including a "collar"
running from behind the gill covers up to the back just in front of the dorsal fin and a
"mask" which runs up from the lower jaws to the eyes then angles forward around
to the front of the face.
Gg. gymnogenys, like Gg. balzanii, is a polygynous, delayed, maternal
mouthbrooder. The other members of the gymnogenys complex are Gg. labiatus,
Gg. lacustris, and, quite notably, Gg. australis which, although known
widely by name in the hobby, is unlikely to have ever been kept in aquaria.
Biotodoma Eigeomann & Kennedy 1903
The genus name Biotodoma means living-home and refers to Eigeomann &
Kennedy's mistaken belief that these cichlids are mouthbrooders. In fact, however,
Biotodoma species are substrate spawners and only move their fry from place to place
in their mouths, in the manner of other substrate spawners.
The type species of Biotodoma is B. cupido and is sympatric with the
other nominal species of the genus, B. wavrini. Both of these species are quite
unimpressive as juveniles, but adults possess wonderful rose and coppery colors and blue
lines (males) or spots (females) on their faces.
A third, and as yet undescribed, species of Biotodoma has found its way into
the aquarium hobby. B. sp. "Red Fin Tocantins" is a strikingly colored
fish with a lyrate caudal fin and double "pin-stripe" lines on their flanks.
They can be sexed as the other species in the genus.
Retroculus Eigenmann & Bray 1894
The species of Retroculus are some of the most highly specialized cichlids in
the world. Not only are they Geophagines (with all the associated adaptations for
sifting), but they are also specialized rapids dwellers with the elongated bodies and
reduced swim-bladders that are favored in fast flowing aquatic environments.
Three species of Retroculus are recognized, R. lapidifer, R.
septentrionalis, and R. xinguensis. Leibel reports that, while R.
lapidifer comes from moderately cool and highly oxygenated waters in the wild, they
are hardy fish in more typical tropical aquariums.10
Cardwell reports that R. xinguensis is found over a sand substrate and will
bury itself quickly in the sand to avoid perceived dangers (such as collectors with cast
nets).
Acarichthys Eigenmann 1912
Acarichthys is a monotypic genus with the single species being A.
heckelii. This species is not a true Geophagine as it lacks all of the specialized
gill morphology that characterizes that group. I have included it here, however, as it
resembles the true Geophagines in most other respects and "fits" well in an
article about them. A. heckelii is most definitely a sifting species and
Steindachner described it as Geophagus thayeri in 1875. (This was over 25 years
after the species was first described, however, so G. thayeri is an invalid
junior synonym.)
A. heckelii do differ from all the true Geophagines (and all other cichlids
for that matter) in the manner of their breeding, however. They are substratum spawners,
but the females dig extensive tunnel networks in the substrate and actively court the
males from these tunnels and the surrounding territory.
Guianacara Kullander & Nijssen 1989
The species of Guianacara are also not true Geophagines but, like Acarichthys,
resemble them in most ways. They also have many similarities to Aequidens species
and other Acaras so Leibel often includes them in articles on Acaras. In fact, Guianacara
geayi was originally described as Acara geayi and known for a long time as
Aequidens geayi.
Kullander briefly and provisionally assigned this species to Acarichthys
(apparently, mostly because he didn't know what else to do with it) but, after further
research, he and Nijssen decided to create a genus just for geayi and its sister
species, which they described at the same time.
From spawning reports and photographs it appears that more than one species of
Guianacara has been found in the hobby under the name geayi, although the
true G. geayi does not appear to have been among them. I have found both G.
owroewefi and G. sphenozona in store tanks in the midwest. These fish, like
most species in the genus, are distinguished by subtle differences in the black markings.
The easily distinguished G. sp. "Red Cheek" has also recently found
its way into the hobby and been spawned by dedicated aquarists. All Guianacara
species look a bit like overgrown, golden rams with black markings.
Papiliochromis Kullander 1977
Papiliochromis ramerezi, the popular Ram of the aquarium hobby, is a true Geophagine,
despite its small size. This fact is reflected in the other generic name sometimes applied
to this fish, Microgeophagus Papiliochromis, for its part, means (roughly)
"butterfly cichlid" and refers to the other common name for Rams. (According to
the somewhat twisted rules of zoological nomenclature, Microgeophagus is
currently the proper generic name for this species, although it was never properly erected
and, I'm confident, will eventually be discarded in favor of Papiliochromis.)
A second species in this genus was recently recognised. The Bolivian Ram, P.
altispinosa, is a higher-bodied fish with a lyrate caudal fin.
Apistogramma Regan 1913
Also among the true Geophagines are the species of Apistogramma. This genus
includes the smallest of all new world cichlids and includes at least sixty species, many
of which remain to be described. There are at least a half-dozen groups or complexes
within Apistogramma, although not all species fall neatly into one of the
divisions.
Apistogramma species differ from most other Geophagines in a number of
respects, mostly having to do with reproduction. They are also more elongate than most
Geophagines.
Apistogramma species are harem-polygynous, cave-spawning cichlids. A male will
guard a territory in which several females breed and the care of the eggs and fry is
exclusively the female's domain.
Like most cave-spawning cichlids, territorial males and females are easily
distinguished. Territorial males are larger, more colorful, and have more impressive
finnage. Outside of spawning, females are generally light grayish or tannish with muted
black markings. When spawning or spawn tending, on the other hand, females acquire a
bright golden-yellow base color with dark black markings.
Quite interestingly, there are non-territorial males which still manage to reproduce.
These males look remarkably like females, both to aquarists and to territorial Apistogramma
males. By fooling the territorial male, these "sneaker males" are able to slip
into the caves of ready females within the territorial male's dominion and fertilize some
or all of the eggs before the dominant male has the chance to.
"Sneaker males" only occur in situations where some males are unable to
establish territories of their own and will become territorial males themselves, with the
accompanying impressive colors and finnage, if a space opens up in the region's hierarchy.
Apistograminoides Mcinken 1965
The final genus of Geophagine is the monotypic Apistogrammoides. As implied by
the name, Apistogrammoides is closely related to Apistogramma. It
differs primarily by the number of hard rays in the anal fin. In Apistogramma,
three to four rays are typical. In Apistogrammoides, the range is seven to nine.
The single species in Apistogrammoides is A. pucallpaensis, which is
named for its collecting site of Pucaflpa, Peru. A. pucalipaensis can be treated
by aquarists as they would species of Apistogramma.
Keeping
Geophagines span the spectrum of cichlid difficulty, from hardy, easily kept and bred
species ideal for beginners, to demanding and difficult fish that are more challenging
than discus. In general, all members of a genus can be kept in the same approximate
conditions, but for breeding it may be necessary to know the precise species that one is
dealing with (see Breeding below).
Water Parameters
Geophagines can be found in all of the water types in South America (see Part One of
this series). Species of Guianacara (which come from the rivers of the Guianan
shield), the Hump-head Earth-eaters (from Panama and the northern flowing rivers of
Columbia), and the Brasiliensoids (from the coastal rivers of Brazil) all hail
from moderately hard, and relatively neutral waters. These fish can be comfortably kept
and spawned in a wide range of pH and hardness values, as long as extremes are avoided.
Most other Geophagines come from more cosmopolitan genera, so species specific
information is needed for determining ideal water conditions for them. If this information
is not available, however, they can be kept in soft to moderately hard (DH 5-10), slightly
acid to neutral (pH 6-7) water without trouble. For breeding, however, some species (such
as daemon complex of Satanoperca and many, though certainly not all,
Apistogramma) need water that is very soft (nearly unmeasurable hardness) and acid
(pH around 4). These conditions can stress members of other species, however, so it is
imperative that the aquarist know the breeding requirements of the species that he or she
is dealing with before attempting to provide these sorts of extreme conditions.
Interestingly, it has been shown that some Geophagines that have evolved to handle low
pH levels in nature actually grow faster in water with a more neutral pH.11 This is
apparently because they only encounter low pH during the dry season as water levels fall
and acids become concentrated. A reduced growth rate during the dry season means that
these fish require less food to survive during these lean times. When the rains increase
again and food becomes more abundant (and the pH rises), the cichlids' growth accelerates.
Almost all Geophagines need warm water in the range of 75-80°F to prosper. Some
species, like those of Biotodoma, only grow well in even warmer temperatures --
up to 85°F. The notable exceptions to this rule are the brasiliensoids which
will live and breed in the home aquarium at 70°F. Also, species of Gymnogeophagus
may be kept around 70°F, although they generally cannot be induced to spawn at this
temperature.
All Geophagines do best in clean water and regular partial water changes are essential
to maintaining healthy fish. Especially vulnerable to poor water quality are species of Satanoperca,
Geophagus, and Biotodoma.
Most Geophagines come from relatively calm waters. The exceptions here are the
Hump-head Earth-eaters which come from flowing waters, and species of Retroculus
which come from swiftly flowing areas including rapids.
Tank Set-Up
Geophagines are, first and foremost, sifters. They will spend most of their time during
the lighted hours of the day taking up mouthfuls of sand and attempting to separate out
any food particles that they can find. For this reason, the choice of substrate for a
sifter tank should not be taken lightly. Large pebbles or gravel may become lodged in a
Geophagine's mouth and cause injury or death. Sharp gravel is also, quite obviously, also
a poor choice. A lack of substrate will not injure these cichlids but will frustrate their
natural instincts and deprive the keeper of witnessing their natural behavior.
The best choice for substrate in a Geophagine tank is rounded silica sand. This inert
sand will pose no threat to the fish, will not effect water parameters, and has a natural
look to it. Of course, other choices of coarse sand or fine gravel may also prove
adequate. Obviously, these fine substrates exclude the use of undergravel filtration.
Large rocks, driftwood, and submerged roots make effective territorial boundaries and
spawning sites while retaining a natural look to the aquarium. Appropriately sized flower
pots and pieces of PVC piping make practical hiding places and spawning caves if
appearance is not a major concern.
The dwarf Geophagines will do quite well in planted tanks and will pose no treat to the
plants, but larger species may uproot or tear apart live plants. This is not true for all
large Geophagines, however, and sturdy, well rooted, potted plants are compatible with
species of Geophagus, Satanoperca, and Biotodoma, and with
Gymnogeophagus balzanii.
Food
Geophagines, despite their natural habits, will quickly learn to take food from the
surface of the water in captivity. However, many species remain clumsy at this unnatural
feeding situation and, if kept with more efficient surface feeders, should be given
sinking foods to ensure that they receive their fair share.
Prepared foods can make up the bulk of the sifter's diet for keeping. These should be
supplemented with frozen and/or live foods such as blood worms, brine shrimp, and glass
worms. For breeding purposes, the balance should shift to frozen and live foods for the
bulk of the diet and feedings should be heavy (and accompanied by many partial water
changes to maintain water quality). The dwarf Geophagines may also be conditioned on baby
brine shrimp.
Greens are unlikely to be eaten by most sifters, although peas may be accepted. Feeder
fish are likely to grow and prosper in a Geophagine tank, as these fish are quite awkward
piscivores. They will, however, sift hiding fry from the substrate if they get the chance.
Tank Mates
Most Geophagines are peaceful cichlids and make good community tank residents. The most
aggressive Geophagines are the brasiliensoids, which can hold their own with all
but the most aggressive Cichlasomines. The recommendations for tank mates listed below do
not necessarily work well for brasiliensoids, which should be kept with the same
types of tank mates as large Acaras (see Part One). At the other end of the scale
are Satanoperca species which can be housed with Angels and small Tetras without
problem. As long as a Tetra is fast enough to avoid an Angelfish, it is more than quick
enough to escape the attentions of Satanoperca.
Despite their peaceful nature, most sifters are able to hold their own with somewhat
more aggressive cichlids, but do their best if kept with other peaceful fish. Good choices
for tank mates for most large Geophagines include all Acaras except Green Terrors
(as discussed in Part One of this series), High-bodied South American Cichlasoniines
(covered in Part Two of this series), Tetras and other peaceful characins, loracariids of
all sizes, carichthyids (Corydoras and related genera), and small pimelodids.
Note, however, that while all of these fish are compatible with large Geophagines, they
are not all compatible with one another.
Dwarf Geophagines can be kept with dwarf Acaras and the remaining dwarf South American
cichlids (which will be covered in the final part of this series), the more peaceful (and
non-piscivorous) High-bodied South American Cichlasomines (e.g. Angels, Discus, and
Festivums), Tetras, loracariids, and callichthyids.
Dither fish, such as Tetras, are highly recommended for tanks containing dwarf
Geophagines, and may also help to calm large Geophagines and encourage breeding.
Breeding
Geophagines show a diversity of breeding strategies and behaviors unmatched in any
other group of New World cichlids. Further, as mentioned above, breeding strategies and
behaviors may vary from species to species within a genus. Finally, the species within a
genus may come from different water types. For these reasons it is important to know the
particular species one is dealing with in order to be prepared to spawn them.
The hardy brasiliensoids, like many popular Cichlasomines, are easily spawned and make
good beginners fish. The recently popular 'G.' steindachneri likewise makes a
good beginners cichlid. It is easily kept and will spawn given a diet of only prepared
foods. In fact, perhaps the only way to prevent a pair of these cichlids from spawning,
short of separating the male and the female, is to reduce the tank temperature below
70°F. The fry of 'G.' steindachneri are large enough to take finely crushed
flake food as a first meal, although they will do better on a diet of newly hatched brine
shrimp.
At the other end of the scale, only one species of the daemon complex of Satanoperca
has ever been successfully spawned in captivity (S. daemon itself) and this only
once. The cichlid enthusiast looking for a breeding challenge should move right past the
comparatively easy Discus to the members of the daemon complex, but only if he or
she is truly prepared. These fish do NOT make good beginners cichlids. Not only are they
extremely difficult to spawn, but many very experienced aquarists have trouble just
maintaining them in captivity.
Other Geophagines which are challenging to breed include Acarichthys heckelii, the
other Satanoperca species (the juruparoids), Biotodoma species, and many
species of Apistogramma. One of the keys to spawning many of these species is
patience. Many of the large Geophagines grow quite slowly and are not mature until they
are a few years old. Dwarf species, on the other hand, mature quite rapidly (they are
essentially annual fish) and may spawn at only two months post-hatching.
Another factor that makes spawning many Geophagines difficult is sexing the adult fish.
As is the general rule with cichlids, when parental duties are shared equally by both
parents, the fish as essentially sexually isomorphic. Species of Geophagus and Satanoperca,
and Acarichthys heckelii are particularly hard to sex. Biotodoma species
can be sexed by their facial markings as noted under the description of that genus. Gymnogeophagus,
Guianacara, Papiliochromis, and the brasiliensoids can be sexed by size
differences (males are larger) and cranial profile (males have steeper faces). Territorial
males of Apistogramma can be recognized at a glance, but subdominant males may be
difficult to distinguish from females (see Apistogramma, above). Finally,
Hump-head Earth-eaters can be sexed easily by the humps of the males and by males' more
impressive coloration.
Blackwater Geophagine species are especially difficult to spawn as they need very soft
water in order for their eggs to be fertilized (in hard water the eggs close up before the
sperm can penetrate) and a low pH to trigger spawning and allow for a reasonable sex ratio
in the fry.
Finally, because most Geophagines are so peaceful, a crowded tank or aggressive take
mates will inhibit spawning behavior.
Most Geophagines lay their eggs on flat surfaces, such as smooth rocks, although the
immediate mouthbrooders are less likely to be fussy about their spawning site than delayed
mouthbrooders or substrate spawners. As noted above, S. daemon lays its eggs in a
depression it digs in the substrate. Some Geophagines, notably species of Satanoperca,
will cover their eggs after spawning, so do not be alarmed if sand sits where the eggs
"should" be.
Apistogramma and Guinacara species and Acarichthys heckelii
are sheltered spawners and appropriate cave or tunnel-like areas should be provided. For A.
heckelii it may help to fill the intended spawning site with sand and allow the
female to excavate it herself.
In all, Geophagines exemplify the degree of diversity that may be found within a single
cichlid lineage, especially in terms of breeding behaviours. The diversity of New World
cichlids, however, is much broader than that of any one lineage. Some of the most unusual
and interesting New World cichlids are yet to be covered. For those, see the next and
final instalment of this series.
1 Axelrods Atlas, for example, shows G.
tacniopareius (p. 355, picture 5), G. brachybranchus (p. 355, pictures 6-8),
G. proxixnus (p. 356, picture 1), and G. altifrons (p. 356, picture 2),
all labeled as G. surinainensis.
2 Leibel, 1990, p. 17.
3 Ibid.
4 Schlesff, 1994, pers. comm.
5 ???, 1994, pers. comm.
6 Leibel, 1994, pers. comm.
7 Hougen, 1993
8 1980a, 1980b, 1981, 1995.
9 According to systematists currently working with
Gynmogeophagus. See Reis, et al., 1992, p. 271.
10 1989, p. 24. 11 See Newman.
Bibliography
Axelrod, Herbert R. and Leonard P. Schultz. 1993. Handbook of Tropical Aquarium
Fishes. T.F.H. Publications, Neptune City, NJ.
Axelrod, Herbert R. "The Golden and Normal Ram: Microgeophogus (Apistogramma)
ramirezi." In Breeding Aquarium Fishes, Book 2, T.F.H. Publications,
Neptune City, NJ, 344-352.
Axelrod, Herbert R, Warren E. Burgess, Neal Pronek, and Jerry G. Walls. 1989. Dr.
Axelrod's Atlas of Freshwater Aquarium Fishes, 3rd Edition. T.F.H. Publications,
Neptune City, NJ.
Burgess, Warren E. 1982. "It's Microgeophagus!" Tropical Fish Hobbyist,
30 (10):7.
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