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The Amphilophus labiatus Species Complex

By , 1982. printer
Published
Paul V Loiselle, 2004

Classification: Species overview, Central and North America.

(This article was originally published in "Frashwater and Marine Aquarium" April 1980, pp. 30-35, 67. It is here reproduced with the permission of author Paul V. Loiselle).

Perhaps the most noteworthy development in the aquarium hobby has been the explosive increase in the number of available cichlid species since the mid 1960's. The large absolute number involved conveys the impression that this phenomenon involves African cichlids exclusively. Such an impression is highly erroneous. Prior to the pivotal year 1964, the rich cichlid fauna of Central America was represented by three commercially available species: Thorichthys meeki, the Firemouth; Archocentrus nigrofasciatus, the Convict Cichlid; and Cichlasoma octofasciatum, the Jack Dempsey. Today, thirty Central American cichlids are available, a ten-fold increase (Editor note: the article was written in 1980, today in 1998 almost a hundred Central American cichlid species are available in the cichlid hobby). Of these twenty seven additional species, five are re-introductions of species earlier imported into Germany or the United States while twenty two are complete newcomers.

Amphilophus labiatus
Amphilophus labiatus
Amphilophus citrinellus
Amphilophus citrinellus
Figure 1: (Top-left) A young male of the red and white piebald morph of the red devil, Amphilophus labiatus. The color pattern of this strain, developed by Beldt's Aquarium. Figure 2: (Top-right) A male of the solid red morph of Amphilophus labiatus. Careful attention to diet has preserved the vivid coloration of this specimen, owned by Jean K. Davies. However, the massive hypertrophy of the lips seen in freshly imported or pond-reared animals is barely hinted at in this red devil. Figure 3: (Bottom-left) A breeding pair of the gold morph of Amphilophus citrinellus. The nuchal hump of the male, the larger of the two fish, is only modestly developed for a sexually active individual. Figure 4: (Bottom-right) This parental female of Amphilophus citrinellus shows the normal banded morph at its most colorful. Were it not for the existance of Oligomelanic morphs of both species, it seems most improbable that either A. labiatus or A. citrinellus would be kept as an aquarium fish. Photographed at Steinhart Aquarium.Photos by Paul V. Loiselle.

Without doubt, the most spectacular of these newcomers were the large, orange-red, thick-lipped cichlids from the Great Lakes of Nicaragua, christened Red Devils. Schofield (1965a) commented on their variability with respect to lip development and body color, a fact also recognized in the vernacular German name for A. labiatus ­ Variabler Buntbarsche -. After consulting several ichthyologists, some with field experience in Nicaragua, Schofield (1965b) tentatively concluded that the large-lipped, orange-red cichlids were Amphilophus labiatus, while the names A. citrinellus and A. erythraeus applied respectively to the yellow and red color forms of another, simple-lipped species. The chief difficulty in correctly identifying these fish stemmed from their remarkable polymorphism, which led, in the context of the limited material available to early workers, to the description of eight nominal species. As pointed out by Miller (1966), by even the most generous appraisal, the total number of published names substantially exceeded the number of biological entities in question. The most recent treatment of this problem (Barlow and Munsey, 1976), concludes on the basis of morphological, behavioral, developmental and ecological evidence that all of the nominal species can be related to the synonymy of two, Amphilophus labiatus (Günther) and A. citrinellus (Günther), while describing a third, previously undiscovered derivative of the latter, A. zaliosus.

These three species were assigned to the subgenus Amphilophus of the then Cichlasoma genus. The fourth member of the quartet I will treat herein, Amphilophus trimaculatus (Günther) is generally referred to the genus Nandopsis. I believe this classification to be in error. Amphilophus trimaculatus lacks the wide, highly protractile jaws of Nandopsis tetracanthus, type species of that genus, and of related Parachormis species such as P. dovii and P. managuense. It also differs completely from the true guapotes with respect to color scheme and sexual dimorphism. In general morphology and behavior A. trimaculatus is virtually indistinguishable from Amphilophus of the A. labiatus group. The same observation can be made of female breeding dress. I consequently believe that A. trimaculatus may more reasonably be regarded as the Pacific slope cognate of the A. labiatus species group. Table 1 presents the scientific and common names of these four species, their dates of importation into the United States, their synonymies and a synopsis of relevant distributional and ecological data.

Aquarists rarely have trouble recognizing A. trimaculatus, whose distinctive color pattern sets it well apart from the other three species in question. Neither are large, wild-caught adults of the other three species easily confused. However, distinguishing juveniles ­ particularly prior to color metamorphosis ­ is difficult under any circumstances. The aquarist, dealing with tank-raised specimens, may also be forgiven occasional perplexity about the identity of his pets. This confusion arises from two sources. First, the enlarged lips of A. labiatus regress quickly under aquarium conditions, while the color of both the Midas Cichlid and the Red Devil tends to fade in captivity. This complicates the identification of even genetically pure stocks.

Amphilophus labiatus
Amphilophus labiatus
Amphilophus citrinellus
Amphilophus citrinellus

Figure 5: (Top-Left) The spindle-shaped body and distinctive lateral banding identify this specimen as Amphilophus zaliosus. In nature, the arrow cichlid is an open water dwelling species, unlike its bottom dwelling relatives. Photographed at Steinhart aquarium. Figure 6: (Top-right) A sexually inactive male Amphilophus trimaculatus. The vivid coloration of the iris has led to this species being christened Red-eyed cichlid by some aquarists. Figure 7: (Bottom-left) A sexually active male Cichlasoma trimaculatum. The derivation of both the scientific name of this species as well as its common name of three-spot cichlid is self-evident from this photograph. Figure 8: (Bottom-right) Parental female Cichlasoma trimaculatum. Note the similarity between the color patterns of parental females of this species and Amphilophus citrinellus. The distinctive pattern of lateral spots is almost completely obscured in the female Three-spot cichlid at this time. Photos by Paul V. Loiselle.

Second, the small numbers of fish initially imported, the confusion over the number of species involved and the great eagerness to spawn them led initially to a great deal of indiscriminate hybridization. While all these cichlids display strong preferences for conspecific mates, they will hybridize in a no-choice situation (Bayliss, 1976). There is even some evidence that introgressive hybridization between A. labiatus and A. citrinellus may have occurred in the past in some of the smaller crater lakes (Barlow, 1976). The progeny of such shotgun marriages are as viable and fertile as the parental species. Thus the first few tank-reared generations were pretty much of a genetic omelette. With the passage of time, these fish have converged phenotypically on A. labiatus. Subsequent commercial importations have consisted exclusively of large-lipped, bright red animals. These obvious A. labiatus have been pond-bred in Florida for nearly ten years, and their offspring have dominated the market during this interval. The original hybrids have been effectively swamped through crossing with such pure A. labiatus.

Today, most tank-reared Red Devils are phenotypically recognizable A. labiatus, though one still encounters the odd individual whose deeper body and blunter snout proclaim the presence of a Midas Cichlid in the rock pile in the recent past!

To return to the problem of telling one species from the other, Table 2 presents several diagnostic features aquarists should find useful in identifying these fish. Additionally, individuals of A. citrinellus are usually deeper bodied than are comparably sized specimens of the other two species. However, this characteristic is influenced by environmental considerations and varies so markedly that it is not a reliable means of differentiating between these cichlids.

Of the four species, A. labiatus is by far the most commercially available, followed by A. trimaculatus. All the A. citrinellus available to hobbyists are ultimately descended from pure stocks maintained by Barlow for research purposes. The same is true for A. zaliosum, which has been released only in small numbers. Its unattractive coloration and belligerence have apparently discouraged efforts to breed it, and I doubt the Arrow Cichlid will ever be available to hobbyists. There are too many more desirable Amphilophus available to make A. zaliosum worth a breeder's while to spawn and rear.

Overhead view of Amphilophus citrinellus and labiatus Figure 9: Overhead view of the upper jaw of Amphilophus labiatus (A) and Amphilophus citrinellus (B). Drawing by Paul V. Loiselle.

The maintenance requirements of this quartet of species are no different from those of other large cichlids. These have been detailed elsewhere (Loiselle, 1978a,b; Mortensen, 1978) and I will not repeat them here. I will emphasize the importance of careful management of the nitrogen cycle to their successful maintenance. A high-speed outside power filter charged with an easily cleaned, reusable medium such as Ehfi-Fix® will remove the particulate waste produced by these gluttonous, messy feeders. 50% to 60% water changes every 7 to 10 days will prevent a dangerous build-up of toxic dissolved metabolites.

Like other large Cichlasoma species, these fish are easily fed. Any of the usual live foods and meaty prepared foods are eagerly taken. If not fed the appropriate pigment precursors, notably b-carotene and canthaxanthin, Midas Cichlids and Red Devils will lose their vivid coloration. Live goldfish, appear to provide one source of these precursor compounds. Most aquarists will find the formula developed by Barlow and Capetto (1974) more convenient and economical.

The extreme belligerence of these cichlids poses the greatest obstacle to their successful management. Though it pains me to say it, all the unfavorable things written about cichlids over the last 75 years are true of this foursome. They are predatory, aggressive and dig prodigiously! Single specimens housed with comparably sized fishes in large (300 l-400 l) aquaria pose the fewest problems. Single specimens housed alone in smaller tanks (100 l-200 l) make highly interesting "pet fish." I add, in passing, that obsessive digging and such quirks as flipping filter intake tubes and heaters onto the floor are more often encountered among such solitary fish. I don't think I am too far off the mark when I ascribe such behavior as response to the acute boredom generated by such a stimulus-poor environment. Serious problems usually arise when several specimens of the same species are housed together alone or in a community tank. Fairly social in nature, all are acutely intolerant of the proximity of conspecifics in captivity. Unmated fish arrange themselves in a linear peck order whose operation results in the emergence of a single survivor, or, if their tank is large enough and both sexes are present, a single pair. Such a pair will live fairly amicably with other large fish until the female is ready to spawn. Survival of their tankmates past this point is purely a function of tank size. If they can move beyond the limits of the pair's territory and stay well away, they are safe. If not, their remaining existence will be brutish, unpleasant and brief. Providing rock caves and breaking the bottom up into several territorial foci will help matters somewhat, but it is sorely tempting fate to try housing a pair of these fish in a community tank of less than 400 l capacity.

Amphilophus labiatus (Günther)
Red Devil Cichlid

Synonyms Cichlasoma froebeli (Agassiz) Cichlasoma lobochilus (Günther) Cichlasoma erythraeum (Günther) Cichlasoma dorsatum (Meek)
Date of importation 1964
Range Lakes Managua and Nicaragua
Habitat Ubiquitous in both lakes but most abundant over rocky bottoms
Feeding pattern Aquatic invertebrates, mainly snails, organic detritus and smaller fish
Polymorphic Polymorphic with respect to both coloration and lip development
Cranial profile and Nuchal hump development Sharply indented cranial profile, moderate nuchal hump
Lip development and shape of the upper jaw seen from above Lips heavily lobed in large wild specimens. A distinct median knob present on the upper lips in tank-reared specimens. Shape of the upper jaw like a Gothic arch when seen from above (Figure 9a)
Rank With Regard to head and snout length: 1 (Longest) 2 (Shortest) 1
Oligomelanic color forms Orange-red to flesh pink; solid white and red, white and black piebald morphs are known
Lateral bar shape in normal morphs Broadest at the center

Amphilophus citrinellus (Günther)
Midas cichlid

Synonyms Cichlasoma basilaris (Gill and Bransford), Cichlasoma granadense (Meek)
Date of importation 1964
Range The Rio San Juan system, including Lakes Managua and Nicaragua and many small crater lakes such as Masaya and Jiloa
Habitat Ubiquitous in the lakes but most abundant over rocky bottoms. Uncommon in riverine biotopes
Feeding pattern Aquatic invertebrates, mainly snails, organic detritus, smaller fish, fish eggs and stranded insects
Polymorphic Most populations polymorphic with respect to coloration only, but L. Masaya fish are also polymorphic with respect to lip development
Cranial profile and Nuchal hump development Moderately indented cranial profile, very large nuchal hump
Lip development and shape of the upper jaw seen from above Simple lips. Shape of the upper jaw like a Roman arch when seen from above
Rank With Regard to head and snout length: 1 (Longest) 3 (Shortest) 3
Oligomelanic color forms Yellow-orange to pale cream. Solid white morph knwon. Yellow and white and yellow. White and black morphs are known
Lateral bar shape in normal morphs No

Amphilophus zaliosus (Barlow)
Arrow cichlid

Synonyms None
Date of importation 1972
Range Endemic to Lake Apoyo, Nicaragua
Habitat Open water, not far below the surface
Feeding pattern Zooplankton, stranded aquatic insects and smaller fish
Polymorphic No
Cranial profile and Nuchal hump development Sharply indented cranial profile, moderate muchal hump
Lip development and shape of the upper jaw seen from above Simple lips. Shape of the upper jaw likea Roman arch when seen from above
Rank With Regard to head and snout length: 1 (Longest) 2 (Shortest) 2
Oligomelanic color forms None known
Lateral bar shape in normal morphs Narrowest at the center

Amphilophus trimaculatus (Günther)
Three-spot cichlid

Synonyms Cichlasoma mojarra (meek) Cichlasoma centrale (Meek) Cichlasoma gordonsmithi (Fowler) Cichlasoma cajali (Alvarez & Gutierrez)
Date of importation 1967
Range Laguna de Coyuca, NW of Acapulco, México, southward basin of the Lempa, El Salvador
Habitat Ubiquitous in both lacustrine and riverine habitats throughout its range
Feeding pattern Aquatic insects, smaller fish, snails, aquatic plants and organic detritus
Polymorphic No

Spawning under community conditions usually proceeds with no complications. Assuming the other residents can survive the experience, this is the least chancy approach to breeding any member of this quartet. Attempts to spawn these cichlids on a single pair basis are more dangerous. Unless appropriate precautions are taken, a badly injured or dead female is more likely to result than a successful pairing. The traditional approach involves separating the prospective pair with a transparent partition, which is removed when the female begins to show an ovipositor. This approach has the best chance of success if the female is 85% the size of her prospective mate (Barlow, 1976; Barlow, Rogers and Capetto, 1978) and the tank is well supplied with hiding places she can enter but the larger male cannot. Providing a scapegoat for the pair's aggressive tendencies in the form of an adjacent tank containing other visible larger fish also facilitates pairing. None of these gambits is a substitute for close surveillance of the prospective partners following removal of the partition. Given the speed with which a male can trash a female, even a ten minute lapse of attention can have serious consequences. If the female is ready to spawn, nothing more violent than reciprocal tail beating and brief bouts of jaw locking should follow. If, on the other hand, the male begins biting and chasing the female in earnest, the two fish should be separated and another effort made to pair them up 24 hours later.

If playing marriage broker is not to one's liking, resort may be had to the incomplete divider method of spawning large substratum spawning cichlids. This approach reduces risk of female injury to nothing and is the method of choice when dealing with single pairs of irreplaceable fish. A screen divider either separates the fish or a glass pane stopped 2-3 cm short of tight contact with the tank bottom. On the female side, a vertical spawning substratum is placed no more than 20 cm away from the divider. The fish will go through the spawning sequence even if physically separated. While the distance between the male and the egg mass precludes 100% fertilization of the spawn, even partial success will result in as many fry as most aquarists are likely to want to rear.

Assuming that courtship is accomplished without mayhem, the pair will go through the spawning pattern herein illustrated for A. citrinellus. The grey to olive-tan elliptical eggs measure c. 2.0 mm along their major axis. They hatch in 2 days at 27°C, and the fry are mobile 4 days later. Artemia nauplii, microworms and finely divided prepared foods are eagerly taken. The fry of the three Nicaraguan species also nibble mucus from the flanks of their parents (Noakes and Barlow, 1973). Their growth is rapid, and given that spawns can range from 2000 to 7500 eggs, such attentions can cause serious injury to the usually acquiescent parents, who lack the space to move away from their insistent progeny. Bouts of abrupt fin-flicking are the first indication that this mucus feeding has reached bothersome proportions. If the fry are not separated from their parents at this point their grazing can result in the appearance of ragged fin margins and open sores on the flanks.

Amphilophus citrinellus
Amphilophus citrinellus
Amphilophus citrinellus
Amphilophus citrinellus

Figure 10-12: (Top-Left) Show the pre-spawning interaction between a pair of Amphilophus citrinellus. The female (foreground) shows a prominent ovipositor, a sure sign of inminent spawning. The spawning secuence, Figure 13: (TOP Left and right), Amphilophus citrinellus spawn on a vertical surface, preferably within an enclosed space. A 25 cm. clay flowerpot, cut lengthwise in half, is an excellent spawning site for the Midas cichlid and the other members of this group. The female initiates spawning by making several "dry runs" over the spawning site, followed closely by her consort. With the appearance of the first eggs, spawning begins in earnest. At this conclusion, the female fans and mouths the eggs, while the male keeps intruders at a distance.

Amphilophus citrinellus
Amphilophus citrinellus
Amphilophus citrinellus
Amphilophus citrinellus

Figure 14: (TOP-Left) The yolk-sac fry of A. citrinellus, like those of other substratum spawning cichlids, have paired adhesive head glands that permit them to adhere to the bottom and to one another even in the face of a fairly strong current. Figure l5: (TOP-Right) The immobile fry are usually moved several times to pre-dug pits before they become free swimming. The fish illustrated is the female, but both parents actually share this task. Figure 16: (ABOVE-Left) This swarm of free swimming fry represent about 25% of the total spawn produced by this pair. The remarkable fecundity of these large cichlids is an adaptation to the intense predation pressure faced by these species in nature. It makes the sibling cannibalism that usually begins about four weeks post-spawning seem a blessing to most aquarists rather than a curse. Figure 17: (ABOVE-Right) The hobbyist imprudent enough to stick his hand into the midst of this touching family tableau would most likely have it well bitten for his pains! These robust mojarras have little difficulty drawing blood in their attacks. Despite the vigilance of their formidable protectors, fewer than l0% of the fry produced by a single spanning survive to the age of a month in nature. Photos by Paul V. Loiselle.

By their sixth week, the fry measure from 10.0 mm to 15.0 mm TL. Unless some effort is made to sort the fry, this size disparity will lead to sibling cannibalism by the beginning of the second month. As very few aquarists have the space ­ or motivation ­ to rear several thousand cichlid fry of a species with very limited commercial possibilities, this tendency to dine within the family is usually considered the manifestation of a beneficent Providence rather than a calamity.

The species-typical black markings of all four species develop at c. 3.0 cm TL. The young of Red Devils and Midas Cichlids, unlike those of other oligomelanic cichlid morphs, are not depigmented at birth. Age at color metamorphosis in A. labiatus and A. citrinellus is variable. Shut-down of melanin synthesis in nature is delayed until the fish are 10.0-15.0 cm Sl, too large to be vulnerable to most of their predators. Unconscious selection for precocious metamorphosis in A. labiatus has resulted in strains in which some individuals in a spawn begin to change color as early as three months of age, while virtually all the young have metamorphosed by 18 months of age. The Midas Cichlid seems a bit slower in this regard. Young reared in Barlow's lab rarely begin metamorphosis earlier than their seventh month, and some individuals may wait two or three years before undergoing color change. When one considers that these fish can begin spawning at 9-10 months of age, it is perfectly possible for an individual to have bred several times as a normally colored fish before developing an oligomelanic color pattern. This variability in color metamorphosis has complicated the study of how oligomelanism is inherited in these cichlids. According to Barlow (pers. comm.) yellow A. citrinellus do breed true, all the offspring of oligomelanic parents ultimately turning yellow. The same holds true of A. labiatus (Macrum, pers. comm.).

The first Red Devils imported commanded asking retail prices of up to $500 each. Yet neither A. labiatus nor A. citrinellus has managed to attain the popularity or commercial importance of their African cousins. The economics of mass-producing a fish that must be withheld from the market until color metamorphosis makes it saleable are a partial explanation. But I suspect the problems of managing such large, aggressive cichlids are chiefly responsible for lack of interest in either polymorphic species or in A. trimaculatus. This is unfortunate, for these large Heroine species display a level of interactive behavior with few equals among fishes, and are both hardy and handsomely colored in the bargain. So if you are growing bored with an endless succession of Malawian mouthbrooders and want to take on a cichlid whose management is a real challenge to your abilities as an aquarist, give one of these mojarras a try. They may exasperate, frustrate and infuriate you by turns, but I can guarantee that they will never, ever bore you!

References

  • Barlow, G.W. and R. Cappetto. 1974. A food for all pisces. Buntharsche Bull. (45): 18-21.
  • Baylis, J.R. 1976. A quantitative study of long-term courtship. I. Ethological isolation between sympatric populations of the Midas cichlid Cichlasoma citrinellum and the arrow cichlid, C. zaliosum. Behavioar 59: 59-69.
  • Loiselle, P.V. 1978a. Two living treasures from the land of the Mayans. F.A.M.A. 1(10): 15-17 est seq.
  • Schofield, D. 1965a. The Red Devils. Aquar. J. 36(1): 5-13.
  • Schofield, D. 1965b. The (Red) devil you say! Aquar. J. 36(7): 322-326 et. seq. 67

References (5):

Citation

Loiselle, Paul V. (November 07, 1998). "The Amphilophus labiatus Species Complex". Cichlid Room Companion. Retrieved on November 24, 2017, from: https://www.cichlidae.com/article.php?id=106.