North America?
I'm amazed by the frequency with which I hear aquarists referring to fish such as Jack Dempseys or Firemouths as "South American cichlids." In fact, they are NOT South American cichlids, they are NORTH American cichlids.1 Why so many aquarists are not able or willing to note this difference is a mystery to me. I rarely hear aquarists confusing the cichlids of Lakes Malawi and Tanganyika -- two rather obscure bodies of water that few non-aquarists in the United States have ever even heard of and which are located very near one another in eastern Africa. Yet I hear aquansts incorrectly stating the CONTINENT of origin of many New World cichlids with great frequency. Do these people really believe that North America only extends south to the U.S./Mexican border,2 or do these aquarists simply fail to understand that the differences between North and South America are every bit as important as the dfferences between Lakes Malawi and Tanganyika when it comes to keeping and understanding cichlids?
Just as Tanganyika has a diverse assemblage of cichlid lineages (Tilapiine, Tropheine, Lamprologine, Haplochromine, etc.) and Malawi is dominated by a single lineage (Haplochromine), South America has a diverse assemblage of cichlid lineages (Chaetobranchine, Cichlasornine, Geophagine, Crenicichline, etc.) and North America is dominated by one lineage (Cichlasomine).
Just as many reproductive strategies are found in the cichlids of Lake Tanganyika (maternal, openly polygamous and monogamous mouthbrooding; monogamous substrate and harem polygamous substrate spawning; biparental etc.) and a single strategy dominates Lake Malawi (maternal, openly polygamous mouthbrooding), many reproductive strategies are found among the cichlids of South America (maternal, openly polygamous and monogamous mouthbrooding; monogamous substrate and harem polygynous substrate spawning; biparental etc.) and a single strategy dominates North America (monogamous substrate spawning).
Just as Tanganyika is home to true dwarf cichlids (the tiny shell dwelling Lamprologines) and true monsters (Boulengerochromis microlepis, said to be the largest cichlid in the world3) and the cichlids of Malawi have a more restricted size range, so too, the cichlids of South America range from true dwarf cichlids (e.g. species of Apistogramma, which will be dealt with next issue) to true monsters (of the genus Cichla, which will be dealt with in part five) and the cichlids of North America fall within a more moderate range.
Comparisons could also be made about percentages of cichlids to non-cichlids, recency of evolution, variety of habitats, etc. but I think the point is made. Very real differences do exist between the cichlids of North and South America and between their respective native environments, and the aquarist who wishes to understand his or her fish will take note of them. Of course, this is not to say that simply recognizing the continent of origin of American cichlids is sufficient, but it is at least a start.
Taxinomical Nightmare
With only a couple of exceptions4 both found in Panama and presumably only recently arrived from South America, all the cichlids of North America, including those native to the Caribbean Islands, are descended from a single lineage - Cichlasomine group A. This much seems clear. Beyond this point, however, the taxonomy of the North Amencan cichlids is extremely confused and badly in need of revision. This fact has been known for nearly one hundred years. (Even as Regan carried out his revision in 1905, he knew it would be inadequate.) Unfortunately, no adequate revision is likely to come soon, even though such a revision has been 'forthcoming" for twenty years now.5
The reason for this sorry state of affairs is that the common ancestry and recent evolution of the North American cichlids makes separating them into discrete genera extremely difficult. (In fact, this difficulty points to the problem inherent any time scientists try to fit the natural world into artificial categories -- these clean divisions, which help us to make sense of what we see, are only approximations of the complex and continuous natural world.)
Nomenclatural Niahtmare
As mentioned in previous installments of this series, Regan, in 1905, used Cichlasoma as a catch-all genus for many American cichlids that could not be easily divided into more meaningfull genera. In 1983, Kullander restricted Cichlasoma to a dozen species of Acaras, leaving many American cichlids "orphaned" (without proper generic placement). In 1984, Loiselle suggested that Heros be used as the catch-all genus for these orphaned cichlids. In 1985, Kullander restricted Heros to just the Severum and its closest relatives.
Since then, most scientists have used 'Cichlasoma' (written just like that; the genus Cichlasoma in single quotation marks) for the orphaned cichlasomines.
In 1993, however, unwilling to live with this reasonable practice, Dr. Warren Burgess and Jerry Walls, in an issue of the aquarium journal Tropical Fish Hobbyist resurrected the genus Herichthys (which was originally erected in 1854 for the Texas Cichlid) as the new catch-all genus for North American cichlids. Fortunately, and not surprisingly, this recommendation has been largely ignored.
To further complicate matters, there are a number of generally recognized, informal "sections" into which the North American 'Cichlasoma' are divided. In general, these were erected as genera but are not currently recognised. When a revision of the North American cichlasomines finally takes place, it is likely that many of these divisions will again be regarded as genera. Until such time, however, the sections are still useful divisions for getting a handle on the North American cichlids. Because they are informal, however, there is no authority as to which species belong in which sections, so disagreement between authors may occur.6
So, following the lead of Kullander, Stiassny, and others (and ignoring Burgess and Walls), and using the section names, we have, for instance, 'Cichlasoma' (Herichthys)' eyanoguttatum for the Texas Cichlid.
This is not the end of the story for North American cichlid nomenclature, however. There are a few currently (though not universally) recognised genera of cichlasomines from North America. These include Neetroplus, Paraneetroplus, Herotilapia, and Petenia. Dr. Robert Rush Miller, in a comment made after a talk at the second International Cichlid Conference (in Orlando in 1991), said that we might as well refer to ALL members of the North American Cichlasomine group A as 'Cichlasoma' until thorough revisions are made. To my knowledge, however, no one has followed his advice, and these other genera are still widely used.
Finally, there have been some attempts in recent years, often by under qualified individuals (aquarists rather than ichthyologists), to resurrect old genera, elevate sections to generic status, or create new genera for the North American 'Cichlasoma', but these have been rather poor attempts in general7 and an authoritative revision of the taxonomy of the entire group is badly needed.
Diversity of North American Cichlids
Simply because the cichlids of North America are (almost) all descended from a single line does not mean that they are all alike. In fact partly because they have not had other cichlid lineages to compete with Cichlasomine group A has diversified to fill more niches in North America than it does in South America. For example, in South America the niche of substrate sifting is filled by the highly specialized Geophagines (which will be covered in Part Four of this series). In North America, free from competition from Geophagines, species of Cichlasomine group A have evolved to fill this niche. Other specialization's have also taken place, and these will be mentioned below.
Divisions
'Cichlasoma' Sections
'Cichlasoma (Thorichthys)' Meek 1904
This is probably the best defined section of 'Cichlasoma'. In fact many authors now use Thorichthys as a genus, believing that its inevitable rise to generic status should be recognized now. The most widely known species of Thorichthys in the hobby is the one named for the person who originally erected the genus Thorichthys -- professor Seth Meek. 'Cichlasoma (Thorichthys)' meeki, the Firemouth Cichlid, needs no introduction to most aquarists. Despite its familiarity, however, it might be worthwhile to say a few words about it and its congeners.
First, while the Firemouth has a reputation for having an "evil disposition"8, all of the species of Thorichthys have far more bark than bite. In fact, the eye-like markings on the gill covers of nearly all Thorichthys (they are lacking in 'C. (Th)' affine), and the impressive frontal displays employed by these fishes, serve to keep them out of real fights which would be likely to bring them harm as they are not truly able combatants.
Second, the long snout, sloping face, and high set eyes of Thorichthys bear more than a coincidental similarity to features of the popular "Jurupairi" (Satanoperca leucosticta, see Part IV). Like this well known sifter, the species of Thorichthys have evolved adaptations for seearching through the substrate for food. For the aquarist, this convergent evolution means that, like the "Jurupari", Thorichthys species should not be housed in aquaria with large or sharp gravel for substrate.
The other ten or so species generally included in Thorichthys, including 'Cichlasoma (Aorichthys)' ellioti, the type species of Meek's genus, all resemble the Firemouth in general appearance, but rather than fire-engine red on the belly and throat, they variously have yellow, pink, rose, or other such colors.
Loiselle9 has pointed out that 'Cichlasoma' saivini shares several (possibly important) characters with the Thorichthys species, but I do not believe that any author has included that species in this section.
'Cichlasoma (Archocentrus)' Gill & Bransford 1877.
This section is relatively well defined by comparison to some of the other sections of 'Cichlasoma'. One potential problem point for this section, however, is that most of the species generally included here resemble each other more closely than they do the type species of the old genus Archocentrus, which is, 'Cichlasoma (Archocentrus)' centrarchus, at least in my opinion.
The Flier Cichlid, 'C.' centrarchus has a very high body and small, protruding mouth; characters that reminded its describers of North American sunfishes of the genus Centrarchus. (The common name Flier Cichlid also refers to a sunfish, Centrarchus macropterus, commonly known as the Flier.) The other species accorded to this section tend to be less laterally compressed and lack the prominently protruding mouth.
There are also differences between 'C.' centrarchus and the other species in this section in size (C. centrarchus is by far the largest species in the section), ray counts ('C.' centrarchus has many more rays in its dorsal and anal fins than the others), and spawning and fry tending (see Breeding, below).
The best known member of this section is the easily kept and spawned 'C.' nigrofasciatum, commonly known as the Convict Cichlid. 'C.' octofasciatum, the popular Jack Dempsey, was at one time included in this section, but most authors (e.g. Heijns, Mayland, Konings, and Stawikowski and Werner) count this species among the Nandopsis (see below) and Conkel suggests that it should be placed its own subgenus.
'Cichlasoma (Herichthys)' Baird & Girard 1854.
Herichthys, as mentioned above, was erected as a genus to contain the Texas Cichlid, 'C. (H.)' cyanqguttatum, the only extant cichlid species native to the United States. Its range extends into Mexico and the remaining members of thiss section (with the possible exception of 'C' bocourti which is questionably included in this section by Conkel), such as 'C. (H.)' carpinte, 'C (H.)' pearsei, and a few undescribed species, are found in Mexico as well. Conkel also includes 'C.' labridens in Herichthys, though most authors (e.g. Konings, Mayland, Heijns, and Stawikowski and Werner) list it as a Nandopsis species (see below).
At most this section probably only contains ten species, including the known but undescribed ones.
'Cichlasoma (Mandopsis)' Gill 1862.
This somewhat confused section was referred to as Parapetenia until 1983 when Kullander pointed out that Nandopsis had priority and was available.10 The type of Gill's genus Nandopsis was specified as Acara adspersa Gunther, 1862, which is now said to be synonymous with 'C.' tetracanthus, a large-mouthed predatory cichlid from the island of Cuba. Another Cuban cichlid, 'C.' ramsdeni is generally said to belong to Arandopsis, but recent photos published in T.F.H. suggest affinities with Amphilophus (see below) instead.
To my eye, the species which most resemble 'C.' tetracanthus are the mainland Guapotes 'C.' managuense, 'C.' motaguense, 'C.' loisellei, 'C' friedrichsthaii, etc., and the Haitian Cichlid 'C.' haitiensis. Also generally included in Nandopsis are strong stouted species such as 'C.' grammodes and 'C.' steindachneri; 'C.' minckleyi which has distinct forms (with different dentition and jaw morphology) and a color pattern that resembles species of Herichthys; species such as 'C.' bartoni, 'C.' beani, and 'C.' labridens which show distinct breeding patterns in which rich black obscures non-breeding blues and greens while the remainder of the colors fade to white; and others. All in all there are some twenty to twenty-five species generally said to belong to this section although I doubt that they will all be included here when revisions are complete.
'Cichlasoma (Theraps)' Gunther 1862.
One of the most confused sections of 'Cichlasoma' is Theraps which may contain up to about thirty species. To put some order to this ungainly group, Werner and Stawikowski in 1987 split these fish into Theraps and a new genus of their own creation - Paratheraps. According to Heijns, these authors gave no characters for their new genus, only for its type species which they newly describe as Paratheraps breidohri.11 (This fish is little known in the hobby, but looks remarkably like a 'C.' bifasciatum color morph with a bluish head.) The lack of characters defining the genus makes using this genus unwise, if not impossible.
In 1993, Conkel tried to go Werner and Stawikowski one better, first splitting Theraps into two groups (with his group 1 similar to their Paratheraps and his group 2 including species of their restricted Theraps), then subdividing the first of his groups again. This apparently was not meant to be a formal split, as was Werner and Stawikowski's, just a useful division for hobbyists, but it was botched rather badly even for its more modest goal. Conkel lists several species as belonging to his Theraps group 1B (on page 136) and then goes on to put their descriptions and pictures in the section covering his group 2. Where does he believe these species belong?
The type species of Gunther's genus Theraps is 'C.' irregulars, an extremely elongated, rheophilic (current-loving), cichlid with a pointed face and an inferior mouth. It hops about goby-like, on the substrate and most likely has a reduced swim bladder. The most similar species seem to be 'C. (Th.)' coeruleum, 'C. (Th.)' gibbiceps, 'C. (Th.)' lentiginosum, 'C. (Th.)' nebuliferum, and a couple of undescribed species, although, according to Heijns, Stawikowski and Werner assign two of these species to Paraneetroplus (see below). None of these fish (including the type species) are common to the hobby, so it is unlikely that the casual aquarist would be familiar with any of them.
The better known species of Theraps are large, high-bodied types like 'C. (Th)' synspilum, 'C. (Th)' bifasciatum, and 'C. (Th.)' maculicauda. According to Heijns, these and several other species should be placed in the new genus Paratheraps, and perhaps if that genus is ever adequately defined they will be.
Finally, 'C.' panamense, 'C.' nicaraguense, and 'C.' tuba are all often regarded as Theraps species, although there is strong descent regarding all three of these species (see Oddballs, below).
'Cichlasoma (Amphilophus)' Agassiz 1858.
This section of fifteen to twenty species is also highly confused. Conkel divides it into two groups, while Bussing and Martin and Stawikowski and Werner (1985) divide it into three. Conkel's group 1 corresponds nicely to Bussing and Martin's "labiatum complex" and Stawikowski and Werner's "labiatum- und macracanthus-Gruppe" although Conkel wavers on the correct placement of 'C. (A.)' macracanthum. This group contains 'C.' labiatum, the type species of Agassiz' genus Amphilophus and one of the hobby's Red Devils, as well as the other Red Devil 'C. (A.)' citrinellum, and two or three other species not well known in the hobby. These are generalized predators and generally quite aggressive fish. Loiselle argues that 'C.' trimaculatum groups quite closely with these species (rather than belonging in Nandopsis where it is more commonly assigned).
Stawikowski and Werner appear to follow the lead of Bussing and Martin in the definition of a 'C.' longimanus group within Amphilophus. Perhaps the species of this group best known in the hobby is 'C.' robertsoni, though it is by no means common in aquarium circles. These fish tend to sift the substratum and have long faces and low-slung mouths to accommodate this habit.
Finally, Bussing and Martin place the remaining species of this section in their "ornatum complex" and Stawikowski and Werner place them in their 'C.' alfari group. These groups both contain 'C.' atfari, but I cannot tell if the two sets of authors agree on any characters to define these groups.
Recognized Genera
Neetroplus Gunther 1866.
Erected by Gunther for his new species Neetroplus nematopus, this genus is currently regarded as monotypic although other species have been placed here by later authors. N. nematopus is an elongate fish with a rounded profile. A small, yet highly aggressive fish, it undergoes a dramatic color reversal during breeding. The light grey body darkens and the single dark bar in the center of the body is replaced by a light bar immediately adjacent to the dark bar's prior position. This color reversal is unknown in other species.
Paraneetroplus Regan 1905.
The only fish widely regarded as a member of this genus is Paraneetroplus bulleri, the type species which was described by Regan in 1905 when he erected the genus. According to Heijns, Stawikowski and Werner referred 'C.' gibbiccps and 'C.' nebuliferum to Paraneetroplus in 1987 but one year later Aligayer argued for their placement in Theraps. The elongated body of P. buileri does remind one somewhat of N. nematopus, as the generic names would suggest, but this species is really more reminiscent of a large Theraps resembling the type species of that section, 'C.' (Th)' irregulare.
Herotila pia Pellearin 1904.
This monotypic genus was erected for a fish related to the genus Heros but resembling Tilapia, Herotilapia multispinoso. As the species name suggests, this fish has elevated spine counts in its dorsal and anal fins. These counts overlap with the range for 'C. (Archocentrus)' centrarchus, the type species of that section. These fish also resemble one another in details of their reproductive behavior (see Breeding, below). In fact, these species were confused with one another in the hobby at one time, although that problem has since been resolved. Now, the well-known Rainbow Cichlid, as H. multispinosa is popularly known as a common "beginner's cichlid."
Petenia Gunther 1862.
The final recognized genus of North American Cichlasomine group 1 is Petenia. Another monotypic genus, the only species is P. spiendida, known popularly as the Bay Snook. (The Oligomelanic morph of this species is known as the Red Bay Snook.) An efficient piscivore, the section Parapetenia (now Nandopsis) was named in reference to this genus. This is perhaps the most laterally compressed cichlid in North America.
Oddballs
'Cichlasoma' panamense (Meek Hildebrand 1913).
As mentioned above, there is great contention as to the correct placement of this species. Often regarded as the smallest Theraps species (growing to no more than 5" SL), it is a species which defies easy description. The teeth of adults resemble those of Neetroplus nematopus, and for that reason it was assigned to Neetroplus by Meek and Hildebrand when it was described. It is now believed that the shape of the teeth in the type material is the result of the environment, rather than genetics. Conkel reports that young fry are too small to eat brine shrimp and must be provided with infusoria. If so, then they must have found sufficient microscopic foods in the tank in which I kept them as I added no additional infusoria and they did just fine. I would report that adults seem rather slow-growing and fragil, but this may have been due to poor stock and/or poor keeping on my part, rather than anything inherent to the species.
'Cichldsoma' nicarazuense (Gunther 1864),
Also often regarded as belonging to the Theraps section, 'C.' nicaraguense is one of the most unusual North American cichlids. The body is long like that of the type species of Theraps, but its head is round and blunt. Its mouth is low-slung and dispite a lack of other noticeable modifications for sifting, that is exactly how this species feeds as thoroughly explained by Loisellel2.
'C.' nicaraguense has a unique sexual dimorphism -- mature females have a dark line running from their gill covers back to their caudal fins, whereas males have a single dark spot in the middle of the flanks instead. Juveniles sport both the stripe and the spot at once.
Also unique to 'C.' nicaraguense among the North American cichlasomiines is the fact that its eggs are non-adhesive. Further, the eggs are usually laid in a trough dug between two rocks, rather than on a stone like true open spawners or in a cave like true shelter spawners.
With these and other unique traits in combination, I cannot help but believe that an adequate revision of 'Cichlasoma' will land 'C.' nicaraguense in its own genus.
'Cichlasoma' tuba Meek 1912 and 'Cichlasoma' sieboldii Kner & Steindachner 1864.
Authors disagree as to whether 'C.' tuba belongs in Theraps or Tomocichia. This later genus was erected in 1908 by Regan to hold his Tomocichia underwoodi which he had originally described as Herichthys underwoodi in 1906. Unfortunately, Regan apparently included material from two separate species in his description. One of these species was 'C.' sieboldii which had been previously described (in 1864). The other has since been described as Cichlasoma tuba by Meek. I am not sufficiently familiar with either 'C.' tuba or 'C.' sieboldii to argue about the correct placement of either species and there seems to be no real consensus in the literature.
South American Pacific coast species.
A few species of Cichlasoma group I occur west of the Andes in South America. These fish are more closely related to the cichlasomines of North Amenca than they are to the other cichlasomines of South America. Further, their habitats more greatly resemble those in which the North American cichlids are found than they do the habitats found east of the Andes. For that reason I have chosen to include them in this article.
The only species of these Pacific coast cichlids found regularly in the hobby is 'C.' festae, often called the Red Terror. The common name refers to the intense red body and head color (which is striped with blue on the flanks) and to the fact that it is found alongside the well-known Green Terror, Aequidens' rivulatus (covered in Part One of this series). 'C.' festae is said by Staeck and Linke and by Konings to be a Nandopsis species, though it hardly resembles 'C.' tetracanthus. Loiselle (1985) places it in Amphilophus and I tend to agree with this placement.
Less well-known in the hobby are 'C.' atromaculatum and 'C.' ornatum which resemble 'C.' festae in overall shape (though not in color) and likely belong in Amphilophus as well. (Note that the fish that Loiselle referred to as 'C.' ornatum13 is not the fish pictured under this name by later authors, for example Stawikowski and Werner, 1988, p. 71.)
Keeping
Water Parameters
The water parameters where the North American cichlids are found vary every bit as much as those in the habitats of South American cichlids, but the ranges are not the same. The extremely soft and acidic blackwaters of South America are not present in North America, though some tea-colored, soft, and acid waters are indeed found in North America. At the other end of the scale, there are waters much harder and more alkaline in North America than in the south. Collectors have reported finding "rocks" in cichlid streams which could be broken open to reveal a bit of wood inside. These "rocks" were formed when the bit of wood fell in the water and minerals in the water accumulated around it as it rolled downstream.
North American cichlids and those from the Pacific coast of South America can be found in cooler waters than the high-bodied Acaras covered last issue. Often times these cooler waters are found in mountainous regions, though the cichlids are rarely found above 1500 feet where the water is below about 70 degrees.
Finally, unlike the species covered in the previous two installments which come from slower, calmer waters, many of the cichlids covered in this part come from more active waters and even swift-flowing streams. These latter are called corrientera or correntos by people who live where these species are found.14 Both names refer to the cichlids' preference for currents rather than calm areas. These fish require a high level of dissolved oxygen in their water.
These factors tend to come in combination, so the aquarist should give rheophilous cichlids (which can be distinguished by their elongated bodies and hopping movements) cooler, harder, more alkaline, more highly oxygenated, moving water, and species from calmer habitats (such as the sifters) warmer, softer, more neutral water with less turbulence. In general, however, precise values for water parameters are generally not as important for the cichlids covered here as they are for the High-bodied Acaras. This is because the North American cichlids come from bodies of water that are much smaller and vary to a much greater extent than the massive rivers of South America.
Tank Set-Up
Rocks and drift-wood should form the bulk of the tank decorations for these cichlids. Plants will be eaten by many of them (especially the high-bodied Theraps species), and ripped up by most of the rest. Even sturdy plastics plants, tied securely in place, will not stand up to the attentions of most of these fish for long.
Many of these fish are sheltered spawners (see Breeding, below), so flower-pots (broken or whole) or pieces of PVC piping may be added as "caves." Such caves also make good hiding places for fish being harassed although a better long term solution is to remove the victim to another tank.
It is probably also a good idea to place heaters and filter parts (such as intake tubes for outside power filters) out of reach of the larger of these species. A Plexiglas screen or a piece of "egg-crate" light grating in one corner of the tank to shield these items from abuse works well.
Food
Many of the cichlid species from North America have feeding specializations of which the aquarist should be aware. A staple diet of food sticks or pellets should be supplemented differently depending on the species being kept.
Petenia spiendida and the species of the Nandopsis section are predators; fish make up a large part of their natural diet and frozen chopped fish make a good dietary supplement. Feeder fish (such as goldfish, guppies, or minnows) should be avoided as they carry parasites and diseases which may infect the consuming cichlid.
The high-bodied Theraps species (which Werner and Stawikowski dubbed Paratheraps) consume a good amount of aquatic vegetation in nature and should receive leafy green vegetables and peas in the aquarium.
The more elongated Theraps species, Neetroplus nematopus, as well as some of the species which defy easy classification (such as 'C' panamense, 'C.' tuba, and 'C' sieboldii) are aufwuchs and/or algae feeders and their diets should resemble that given to Malawian mbuna.
Tank Mates
The North American cichasomines range in temperament from the bluffing Thorichthys to the extremely aggressive species that partially make up both Amphilophus and Nandopsis. Despite its small size, Neetroplus nematopus is as aggressive as they come. Besides other cichlids closely matched for aggression, these cichlids can all be kept with loricariids (sucker-mouthed catfishes) of appropriate size, assuming that the catfish have an abundance of spines to fend off the more aggressive cichlids.
Herotilapia multispinosa, 'Cichlasoma' nicaraguense, and the species of the Thorichthys and Archocentrus sections can be kept with moderate sized, fast swimming fish such as tetras of the genus Astyanax or the larger livebearers. Petenia spiendida and many of the species of Nandopsis and Amphilophus, however, would make quick work of such rank mates, and are best kept only with fish able to hold their own in a head-to-head competition.
Breeding
As mentioned at the beginning of this article, the cichlasomines of North America are almost universally monogamous substrate spawners. This means that a pairing will be maintained throughout spawning and fry raising and that extraneous fishes will be badly abused unless they can retreat from the pair's territory.
There is some variation in spawning these cichlids, however. As mentioned above, 'C.' nicaraguense lays non-adhesive eggs. This means that if the eggs are removed from the pair for artificial hatching, something along the lines of the "egg-tumblers" used by Malawi cichlid breeders will be needed to keep water flowing past the eggs without blowing them all over the hatching tank. Further, 'C.' nicaraguense tends to spawn in a semi-sheltered location between two rocks. This behavior falls somewhere between the open spawning of the cichlids covered in the first two parts of this series and the cave spawning exhibited by many dwarf cichlid species. Other North American cichlid species tend more toward one end of this spectrum or the other. Amphilophus, Herichthys, and Theraps species, for instance, tend to spawn more in the open and Archocentrus species go for more sheltered locations. None of these cichlids are highly specialized in this respect, however. Archocenims eggs, for example, are not brightly colored as are the eggs of "true" cave spawners (see Part Four of this series). They also lack "egg-stalks," which many cave spawners' eggs possess.
After hatching, most fry of the North American cichlasomines are moved from pit to pit as were the fry of most of the cichlids covered in Parts One and Two of this series. Herotilapia multispinosa and 'Cichlasoma (Archocentrus)' centrarchus, however, often move their fry to the roots of floating plants, especially if the water has a low dissolved oxygen level.
Finally, the fry of some of the North American cichlids (e.g. members of the 'C.' labiatum complex) feed "Discus-style' from their parents' flanks although none of them require this food if properly fed by the aquarist on newly hatched brine shrimp or micro-worms.
So, while the North American representatives of Cichlasomine group 1 have evolved more diverse breeding strategies than the South American members of this assemblage, for really diverse breeding strategies in the New World Cichlidae one must look to another lineage. That is just what we will do next issue in Part IV: Geophagines, coming in the September/October Aqua News.
1 Hobbyist when not incorrectly claiming that North American cichlids come from South America, usually say that they come from Central America. This is valid, as far as it goes, which isn't very far. Central America is really a socio-political region and fish don't tend to recognize theese sorts of definiations. To drive home the point that these fish are not South American, I have chosen to use the term 'North American cichlid' as most people don't mistake North America for a part of South America.
2 Perhaps some do. One aquarist/author has gone so far as to title his book "Cichlids of North and Central America," as though they were seperate regions.
3 Loiselle, 1985, p. 15.
4 Aequidens coeruleopunctatus covered in Part One, and 'Geophagus' steinduhneri, which will be covered in Part Four.
5 Loiselle, 1991, pers. comm.
6 For example, 'Cichlasoma' panamense (which was described as Neetroplus panamensis) is said by Konings and by Conkel to belong to the Theraps section, by Mayland to belong to Herichthys, and by Staeck and Linke and by Heijns to fall outside any of the currently recognized sections.
7 See Heijns for a partial account of these attempts.
8 Axelrod and Schultz, p. 622.
9 1980b, p22.
10 p. 270.
11 Actually, according to Heijns, 'C.' breidobri was not explicitly designated as the type species of Paratheraps until 1989; yet more evidence of the sloppy work being done in these revisions.
12 1984a
13 1980d and 1981.
14 Conkel 1993, p. 152.
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