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Melanochromis The Beautiful Baddies From Lake Malawi

By , 1979. print format
Published
Paul V. Loiselle, 2004

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Species treated in this document: Melanochromis auratus Melanochromis dialeptos Melanochromis heterochromis Melanochromis loriae Pseudotropheus johannii

(This article was originally published in Freshwater and Marine Aquarium Magazine, Dec 1979; pp. 17-23. It is here reproduced with the permission of author Dr. Paul V. Loiselle).
Melanochromis vermivorous
This adult male Melanochromis vermivorous clearly displays the combination of light lateral stripes on a dark background that characterizes males of this genus. Photo by Paul V. Loiselle.

One of the more gratifying aspects of writing for a nationally circulated aquarium magazine is the feedback generated by your material. It is enormously flattering to receive a phone call from a total stranger who, on the basis of an article he has read, has concluded that you have the answer to his problems in maintaining one cichlid or another, even when the caller's timing suggests a lack of awareness of the time differential between the Atlantic and Pacific coasts! There is even a degree of satisfaction in receiving a letter that points out what the writer considers a mistake on your part. At least he read the article, even if he didn't agree with it! Ego stroking aside, this sort of feedback also allows the writer of a regular column to gauge where areas of intense interest lie and gives him some idea of how he should arrange his research priorities. On the basis of reader response to some of my previous articles, the time seems ripe to do a column on African cichlids. Hence this month's essay will be a review of the distinctive mbuna of the genus Melanochromis.

The genus Melanochromis was established by Dr. Ethelwyn Trewavas in 1935 for an assemblage of mbuna that differed from the genus Pseudotropheus in having fewer and larger teeth on the lower pharyngeal bone. While Trewavas made no explicit mention of the color pattern in her generic definition, she must have been struck by the overall dusky hue of these mbuna, for the name Melanochromis translates literally from the Greek as "black cichlid." Apart from the dental characteristics cited in the generic definition, the fishes included in this genus had little in common. Melanochromis melanopterus and Melanochromis vermivorous, the former the type species of the genus, are long-snouted, rather slender bodied fish with a color pattern that features lateral stripes as its predominant feature. Melanochromis brevis and Melanochromis perspicax are shorter snouted, deep bodied fish with a color pattern that features vertical bars on the flanks. Melanochromis labrosus is a long snouted, deep bodied fish with hypertrophied lips and, unlike other species of the genus, the anterior teeth in the jaws are considerably longer than the other jaw teeth.

Anterior tooth from the jaw of (A) Melanochromis melanopterus and (B) Labidochromis exasperatus. Though both bicuspid, these teeth have little else in common. The sharply pointed, unequally bicuspid dentition argues for this species inclusion within the genus Labidochromis. All other Melanochromis species have the same sort of incisor/like, more or less equally bicuspid dentition as Melanochromis melanopterus. Drawing by Paul V. Loiselle.

Outline and schematic representation of the dentition of the lower pharyngeal bone in (A) Melanochromis melanopterus, (B) Melanochromis auratus and (C) Pseudotropheus zebra. While that of Melanochromis auratus shows the enlarged teeth along the posterior row seen in Melanochromis melanopterus, the degree of crowding of the remaining pharyngeal teeth approaches more closely the condition seen in Pseudotropheus zebra. Drawing by Paul V. Loiselle.

Following the aquaristic debut of the mbuna in 1965, it quickly became apparent that Pseudotropheus auratus was far more similar to M. vermivorous with regard to its behavior and color pattern than it was to any other member of the genus Pseudotropheus. This led my friend and colleague Mike Oliver and I to look at the pharyngeal anatomy of P. auratus. We found that, while the teeth were more crowded than was the case in M. melanopterus, those of the posterior row were definitely enlarged, proclaiming P. auratus a legitimate Melanochromis and thus strongly supporting our suspicion that color pattern was as useful a defining characteristic for this mbuna genus as the nature of the pharyngeal dentition. In 1971, we both had the opportunity to visit the British Museum in London.

During the course of conversations with Dr. Trewavas, we found that her views on the probable generic placement of several nominal Melanochromis had also changed since her erection of the genus. When we examined the types of M. brevis and M. perspicax, it became evident that Mike's initial assessment of their probable affinities was correct and both belonged in the genus Pseudotropheus. None of us were very happy about M. labrosus, but as the unique type was a juvenile, there wasn't a great deal that could be determined about its relationships by a subsequent examination. When additional material became available in 1973, Mike concluded M. labrosus belonged in the genus Haplochromis. This generic placement has since been accepted by aquarists and ichthyologists (Burgess, 1976). While I initially agreed with this assessment, I now have reservations, published at length elsewhere (Loiselle, 1978), whose tenor I would summarize with the prediction that with further study, the generic placement of H. labrosus is very likely to change once again, in a fashion that reflects closer relationship to the mbuna than to such lobe-lipped Malawian Haplochromis as H. euchilus.

With continuing exportation of cichlids from Lake Malawi, the number of known, but undescribed, mbuna species increased dramatically. As early as 1967, several new Melanochromis species had been imported and marketed under erroneous identifications. Since 1973, the situation has been somewhat ameliorated by the publication of a number of papers describing new Melanochromis species (Eccles, 1973; Johnson, 1975; Burgess and Axelrod, 1976; Burgess, 1976), At present, I know of only two Melanochromis species in captivity that remain to be formally described. This situation contrasts most agreeably with that prevailing in the genus Pseudotropheus, where described and undescribed species are about equal in number, or in the genus Labidochromis, where the latter outnumber the former by a conservative three-to-one!

With this brief historical excursion concluded, we can now attempt to define the genus Melanochromis. The cichlids of the genus Melanochromis are mbuna of elongate or spindle-like body form. Their anterior lateral and median outer jaw teeth are equal in size. They are bicuspid in shape and rather incisor-like, the two cusps being of approximately equal length. (Figure 1.) The posterior lateral teeth are enlarged and conical. From three to seven inner tooth rows are present in the jaws. The teeth of the posterior row of the lower pharyngeal bone are somewhat larger than the remaining pharyngeal teeth, whose degree of enlargement and spacing varies somewhat between species. (Figure 2.) The color pattern is based upon lateral stripes rather than vertical bars. In all instances known, males display light stripes on a darker background. In the majority of cases, juveniles and females display the reverse pattern of dark stripes on a lighter background. Table 1 presents a list of the mbuna that correspond to this generic definition, together with prevalent common names for each, modal lengths in captivity, and other relevant information. Table 2 presents a list of mbuna originally described as Melanochromis species under their appropriate generic placement.

The genus may be subdivided into two groups, which differ in cranial morphology and feeding pattern. The species comprising the first group are large mbuna with long, pointed snouts and wide mouths. Their outer jaw teeth are reduced in number, enlarged in size and rather widely spaced. Both the median pharyngeal teeth and those of the posterior row are enlarged and the remaining lower pharyngeal teeth are rather widely spaced. These species, of which M. melanopterus is typical, are invertebrate feeders and opportunistic piscivores. I have observed individuals of M. melanopterus, M. simulans, and M. chipokae wait next to a crevice between two rocks in a mbuna community tank and successfully pick off fry emerging therefrom, while Mike Oliver tells me he has seen solitary M. melanopterus follow parental female Haplochromis of several species and rush in to seize one or two fry once their mother releases them to forage. The so-called Pike Auratus seem to be the morphological and ecological counterparts of the Lamprologines like Lepidiolamprologus elongatus, Neolamprologus tetracanthus and Lamprologus callipterus in Lake Tanganyika.

The second group of Melanochromis comprises small to medium sized mbuna with short, rather rounded snouts, and relatively narrow mouths. Their outer jaw teeth are quite closely spaced and show neither the degree of enlargement nor the reduction in number characteristic of the species comprising the M. melanopterus complex. The teeth of the lower pharyngeal bone are quite crowded and only those of the posterior row show any degree of enlargement. These Melanochromis species appear to feed primarily on aufwuchs, with aquatic invertebrates playing a secondary role. Melanochromis auratus is a typical representative of this group, which comprises those species that are most commonly available to aquarists.

The over.all maintenance requirements of Melanochromis are essentially the same as for other mbuna species (Vide Crout, 1978), but keeping and breeding them successfully is complicated by the fact that they are far more aggressive fishes than the majority of mbuna. Within this context, it is unfortunate that so many novice mbuna keepers elect to make M. auratus one of their first purchases. Though juvenile M. auratus undoubtedly have one of the most dramatically beautiful color patterns of any mbuna species, adult males are, undoubtedly, among the most aggressive of the species available regularly through commercial channels. Male Melanochromis are violently intolerant of the proximity of conspecifics of the same sex. Housing several males together for any length of time is virtually impossible in a tank of less than 400 liter capacity unless the fish are either crowded to a degree rarely seen outside of a wholesale establishment or else kept in all-male groups. Contrary to the case in most other mbuna species, female Melanochromis also quarrel a good deal among themselves. Indeed, females of M. parallelus will indulge in intramural mayhem with every bit as much gusto as their consorts, with essentially the same results. Obviously, the larger the fish, the larger the tank required to dampen out such bellicose tendencies. Hence, the counsel that beginners limit themselves to M. johannii and M. interruptus, whose small adult size make them more manageable in 120-200 liter tanks.

Table 1. The Melanochromis Species to Date (1979) Imported Into The United States
Scientific name Suggested common name Commonly encountered Misidentifications Length Modal Brood Size Comments
Melanochromis melanopterus Species Group

Melanochromis melanopterus Trewawas 1935

Dusky Pike Auratus

-

to 13.0 cm SL

30-50 fry

The male color form known, the blue-dorsal form pictured herein and the more commonly seen form with a broad orange-red margin to the dorsal and caudal fins. An adroit predator of fish the size of a male guppy.

Melanochromis simulans Eccles 1973

Pike auratus

-

to 13.0 cm SL

35-50 fry

Females of this species have the same crisp black and yellow striping as do female melanochromis auratus. Hence the species name simulans. Predatory behavior as for Melanochromis melanopterus.

Melanochromis chipokae Johnson 1975

Black and white pike auratus

-

to 13.0 cm SL

25-40 fry

Predatory behavior as for Melanochromis melanopterus.

Melanochromis loriae Johnson 1975

Golden pike auratus

Melanochromis perspicax

to 13 cm. SL

25-40 fry

Predatory behavior as for Melanochromis melanopterus.

Melanochromis sp. 1

Giant pike auratus

Melanochromis perspicax

to 18 cm. SL

35-50 fry

An adroit predator on fish the size of a feeder goldfish.

Melanochromis sp. 2

Giant black and white pike auratus

Melanochromis perspicax

to 18 cm. SL

No data

An adroit predator on fish the size of a feeder goldfish.

Melanochromis auratus Species Group

Melanochromis auratus (Boulenger 1897)

Auratus

-

to 8.0 cm. SL

35-50 fry

Females differ among themselves with regard to the age at which masculinization of their color pattern occurs. Breeding stock should be selected only from the fry of late-metamorphosing females.

Melanochromis vermivorous Trewawas 1935

Vermivorous, Blue auratus

-

to 10.0 cm. SL

35-50 fry

Female Melanochromis vermivorous are silvery-beige with two narrow black lateral stripes. This drab color pattern, shared by fry, accounts in large measure for the decreasing commercial availability of this species.

Melanochromis parallelus Burgess and Axelrod 1976

Black and white auratus

Pseudotropheus fuscus

to 8.0 cm. SL

25-40 fry

Females behave very aggressively towards one another. They undergo an age-related color change comparable to that seen in female Melanochromis auratus.

Melanochromis johanii (Eccles 1973)

Johanii, Gold auratus

-

to 8.0 cm. SL

25-40 fry

Males display considerable geographic variation with regard to the clarity and extent of the blue lateral stripes. Two female color morphs are known, one beige with olive brown lateral stripes, the other solid yellow-orange. The former has not become established as an aquarium fish. A sexually isomorphic population from the Likoma Island group has been imported. Both sexes are black with blue lateral stripes.

Melanochromis interruptus Johnsosn 1975 Dwarf auratus - to 6.0 cm. SL 25-40 fry Females undergo an age-related color change similar to that seen in female Melanochromis auratus
All mbuna tend to grow larger in captivity than in nature. Thus is is not unsual to encounter the occasional specimen several centimeters larger than the maximun lengths cited in the Table.

Not too surprisingly, in light of the foregoing observations, males of different Melanochromis species don't seem to care for one another's company either. As this difficulty does not extend to females, I suspect its roots lie in the virtually identical color patterns displayed by sexually active males regardless of species. If one wishes to condomicile two Melanochromis species, friction seems to be minimized if one is a member of the long snouted M. melanopterus group and the other is drawn from the short snouted M. auratus group. Outside of periods of sexual activity, other mbuna species are generally ignored, but with its onset, male Melanochromis become vigorously territorial and tolerate no intrusion into their display area by any fish other than ripe conspecific females. It is possible to convey some idea of how important spacious quarters are to the successful management of these beautiful but bellicose mbuna with the observation that a male M. auratus can effectively sequester half of a 200 liter aquarium for his display and spawning site.

Mbuna originally described as Melanochromis sp.
Pseudotropheus brevis (Trewawas 1935)
Pseudotropheus perspicax (Trewawas 1935)
Haplochromis labrosus (Trewawas 1935)
Labidochromis exasperatus (Burgess 1976)

Like all mbuna, Melanochromis are highly polygynous maternal mouthbrooders. Males court potential consorts almost as violently as they defend their territories. Female survival is, therefore, very much a function of giving them enough room to maneuver as well as plenty of shelter. The risk of injury to females is also minimized if the fish are maintained on a harem basis, with two or more females per male. However, given the tendency of females to squabble among themselves, the aquarist must monitor their interactions closely and remove any individual who seems to be suffering from persistent harassment. Actual spawning follows a period of reciprocal circling by the consorting couple that occurs in a sheltered area, often a pit excavated in the center of the male's territory. As in all other mbuna, fertilization of the eggs occurs within the female's mouth. Though males have well developed anal pseudo-ocelli, these are not utilized as egg-dummies. Instead, the female directly mouths the male's vent. Once spawning is over, it is vital that the female have access to shelter, as the male will usually continue to court her vigorously and, in the absence of an appropriate response to his advances, he may injure or kill her.

Ovigerous females are recognizable by an overall muddying of their formerly highly contrasted color pattern of lateral stripes. Interestingly enough, as females grow older this color pattern change may become permanent, their base color darkening so markedly that in such species as M. auratus and M. melanopterus, they become superficially similar enough to males to make distinguishing the two sexes difficult. A tendency towards masculinization of color pattern is known to occur in other mbuna species, such as Pseudotropheus lombardoi and P. minutus. I have even seen very old female H. burtoni with a rudimentary orange humeral patch, an essentially male color characteristic. It is tempting to explain such changes in terms of shifting hormonal balances following upon progressive ovarian depletion. However, this hypothesis does not explain why the phenomenon is almost universal in Melanochromis, only episodic in Pseudotropheus and, to date, unreported in Labidochromis when ovarian depletion is a feature common to the biology of all three genera.

As such masculinized females do not seem at any disadvantage vis-a-vis, their unaltered counterparts with regard to recognition as potential spawning partners by males, the phenomenon also leads one to wonder why natural selection has operated to produce the striking sexual dimorphism that characterizes the genus Melanochromis. Sexual dimorphism in most other mbuna (and indeed in most haplochromid cichlids) usually involves brilliantly colored males and drab females, but among Melanochromis, both sexes are vividly (if very differently) colored. Our present knowledge of the behavioral ecology of these cichlids is so slight that even formulating plausible hypotheses about the function of such sexual color differences is exceedingly difficult. Nevertheless, I think it is worth raising some of these fascinating evolutionary problems both to underline the fact that scientists from many disciplines also find these cichlids intensely interesting objects of study and, hopefully, to arouse in some of this column's younger readers, the curiosity that may someday lead to their elucidation.

Females carry their broods for 21 days at 29°C. Females are usually reliable parents and, given the availability of shelter, rarely have any difficulty carrying successfully in a mbuna community tank, Somewhat surprisingly, given their apparent inability to follow threat with action, parental females become increasingly aggressive towards their tankmates as they approach the end of the incubation period. I suspect that such behavior lessens to some degree the likelihood of unwelcome company when the moment arrives to release their fry. Females will guard their young for several days following their initial release in a nursery tank, taking them into the buccal cavity when danger threatens. I would not be greatly surprised to find that such behavior also occurs in the mbuna community tank as well. If females are isolated during the incubation period, there is little benefit in prolonging the association of mother and fry once the latter are fully mobile. In fact, there is considerable disadvantage to the fry in not separating them from their mother, as her highly protective behavior is likely to interfere with their foraging. Melanochromis fry, like those of other mbuna species, are quite large and are easily reared. They are greedy eaters who have no difficulty with Artemia nauplii or finely divided prepared foods as their initial meal and grow rapidly to the point where they can take adult brine shrimp and standard flake foods. Their coloration will be more intense if they are provided with plenty of vegetable matter in their diet. With heavy feeding and frequent water changes, they will grow to a saleable 3.0 cm TL by the eighth week post-release. Melanochromis interruptus can begin the male color metamorphosis as early as four months post-release at c. 5.0 cm SL. In the other short snouted species, males begin to change color between the sixth and ninth month post-release, while, among the larger long snouted species, the metamorphosis rarely commences before the tenth month post-release. Interestingly enough, males of both M. interruptus and M. johannii are fully capable of fertilizing eggs prior to color metamorphosis, but in the other species of the genus, the onset of male sexual activity appears to wait upon the assumption of adult coloration.

Despite their behavioral shortcomings, most Melanochromis species seem destined to enjoy a permanent place among the ranks of aquarium fish, for they are prolific producers of brightly colored, highly saleable fry. However, the prudent dealer will eschew the temptation to foist a trio of M. auratus or M. chipokae upon inexperienced customers and direct them instead to smaller, less aggressive mbuna species for their initial experience with Malawi cichlids. In so doing, he maximizes the likelihood that a passing fancy will ripen into a full fledged and profitable African cichlid addiction rather than into an equally total aversion. Once a beginner has learned the basics of mbuna keeping with more easily managed species, he will find the many magnificent Melanochromis a stimulating challenge rather than a source of expensive frustration.

Pseudotropheus brevis Haplochromis labrosus

Though originally described as a Melanochromis species. the vertical bars on the flanks of this male Pseudotropheus brevis explain its present generic placement. Photo by Paul V. Loiselle.

Haplochromis labrosus was also originally described as a Melanochromis species. Though one could hardly describe this adult male as brightly colored, the combination of distinctive lip morphology and extroverted personality have won H. labrosus considerable popularity. Photo by Paul V. Loiselle.

male Melanochromis melanopterus female Melanochromis melanopterus

Adult male of the blue dorsal color form of Melanochromis melanopterus, the dorsal fin of the more frequently seen form is sooty and shares with the caudal fin a vivid orange-red margin. Photo by Paul V. Loiselle.

The unproposessing color pattern of this female Melanochromis melanopterus is shared by juveniles as well. Photo by Paul V. Loiselle.

male Melanochromis simulans male Melanochromis sp. 1

Adult male Melanochromis simulans. Females of this species have the same pattern of crisp yellow and black lateral stripes as do those of the better known Melanochromis auratus. Photo by Paul V. Loiselle.

Adult male Melanochromis sp. 1. It is usually offered for sale under the erroneous name Melanochromis perspicax. Females of this largest known Melanochromis species are silvery beige with a faint blueflush on the head and flanks. An indistinct brown midlateral stripe may be present. Photo by Paul V. Loiselle.

male Melanochromis auratus female Melanochromis auratus

The bright yellow pigmentation in the dorsal fin of wild-caught male Melanochromis auratus is only hinted at in this tank-reared specimen. Full intensity will return if captive fish are offered plenty of vegetable diet in their diet. Photo by Paul V. Loiselle.

The splendid coloration of this female Melanochromis auratus belies the general observation that female maternal mouthbrooders are drably colored. Beginning mbuna keepers are advised to resist the temptation to inmediately purchase this stringly marked but very aggressive cichlid. Photo by Paul V. Loiselle.

female Melanochromis parallelus female Melanochromis parallelus

The color pattern of this female Melanochromis parallelus explains the species' color name Black and White Auratus. Photo by Paul V. Loiselle.

The muddying of the color pattern of this female Melanochromis parallelus is as certain an indicator of her ovigerous condition as her bulging throat. Photo by Paul V. Loiselle.

male Melanochromis johanii female Melanochromis johanii

An adult male of the Likoma Island population of Melanochromis johanii. The blue lateral stripes are narrower in males from mainland populations and are frequently interrupted. Photo by Paul V. Loiselle.

The brilliant coloration of this female Melanochromis johanii is much more saturated in newly imported individuals. The silvery olive, brown-striped female morph of this species is virtually unknown in the hobby. Photo by Paul V. Loiselle.

male Melanochromis interruptus female Melanochromis interruptus

The small size at which male Melanochromis interruptus undergo color metamorphosis has earned this species the comon name of Dwarf auratus. This male is only 5.0 cm. SL! Photo by Paul V. Loiselle.

The color pattern of this female Melanochromis interruptus is strongly reminiscent of that of Melanochromis auratus females. However, the dwarf Auratus is a less aggressive species and thus a better choice for the novice mbuna keeper than its larger and better known congener. Photo by Paul V. Loiselle.

Literature Cited

Citation:

Loiselle, Paul V.. (January 14, 1998). "Melanochromis The Beautiful Baddies From Lake Malawi". The Cichlid Room Companion. Retrieved on September 01, 2014, from: http://www.cichlidae.com/article.php?id=80.