|The male Lamena nourissati, the main guardian of eggs and larvae. Photo by Patrick de Rham.|
As most readers of Cichlid News already know, lamena is the native name for a new and beautiful cichlid from Madagascar. First collected by Jean-Claude Nourissat and myself (as juveniles in October, 1991 and then as adults in October, 1992) in the vicinity of Mandritsara, a town in the north central part of the island. At the present time the species is known only from restricted stretches of the Mangarahara River and its rocky, clearwater tributary, the Ambomboa River. Lamena, which translates as "red one" in Malagasy, is a rheophilic species related to the genus Paretroplus. In my opinion, the new species is sufficiently distinct from all known members of Paretroplus to warrant the creation of a new generic taxon; it is anticipated that the formal description of the lamena will appear soon in a scientific journal.
Of the 20 adult specimens collected from the Ambomboa in 1992, I kept four individuals, which I transported in my suitcase to their new home in Lausanne, Switzerland. They were housed in a 130-gallon aquarium with four juvenile Paratilapia polleni, an unidentified Madagascar goby, and for a short time, a few Bedotia geayi and Rheocles species. Water-worn rocks, pieces of driftwood, and reclined flower pots provided many hiding places while simulating the natural environment of the fish. The bottom was covered with a thin layer of sand. Plants consisted of floating hornwort and a few Aponogeton sp. planted in shallow flower pots filled with gravel. Water provided - a mixture of tap and demineralized was soft with a neutral pH, while temperatures were maintained at 24-27°C. Though water changes were frequent and massive, conditions on the whole were very similar to those found in the fish's natural habitat.
Within one or two days of being introduced to the aquarium, three of the lamena had established territories. Of these, the two larger individuals each claimed half the tank, while the third was left with a very restricted space in the middle among a few rocks. These three individuals were beautifully colored in bright orange with two contrasting black bands. Border confrontations were frequent but seldom came to blows. Obviously these individuals were males. The fourth lamena remained a drab brown and did not establish a territory. However within the next few days it had paired off with the male occupying the left half of the tank. Some interaction was observed, and the pair may have spawned, but no eggs were ever observed. Two weeks later the pair bond broke down, and the male became very aggressive, harassing and biting the female who had great difficulty in finding a refuge in spite of the many hiding places provided. In an attempt to relieve some of the pressure on the female, the two other males were removed. The remaining male immediately claimed the entire tank as his territory and intensified his attacks on the female. As her body and fins began to show the effects of his assaults, the female was separated from the male with a glass plate. She quickly revived, her wounds healing in a surprisingly short time. The pair spent most of their time in a face-off at the plate with no signs of hostility. However each time I removed it, the male resumed his persecution. Nonetheless, after a month of short-term trials, I removed the partition permanently as it was preventing good water circulation. The female again went into hiding. Fortunately the aggressiveness of the male diminished with time, and although the female continued to behave in a stressed manner, she did manage to survive. At each feeding she'd quickly grab a chunk of food mainly frozen bloodworms before diving back into her hiding place to eat.
Several months passed in this manner; at the end of May (1993), a change in the behavior of the pair was observed. First, the appetite of the female increased, which led to her emerging from her hiding place each time my presence indicated a feeding was imminent. As a result of her increased food intake, she began to fill out and her abdomen became slightly swollen. As this happened, the aggressiveness of the male declined further. Indeed, his attacks were less frequent and were always preceded by frontal displays, presenting his body laterally with fins erect and quivering. Soon the male's aggressiveness disappeared completely and the pair began patrolling the tank together. After inspecting several sites, the pair concentrated their activities under a large piece of driftwood arched over a flowerpot containing gravel and Aponogeton bulbs. Initially the male took the lead in luring the female to the site, but eventually the roles reversed with the female leading the male to the selected site. At this time the female's oviduct protruded prominently, whereas the male's genital papilla, which had always been slightly visible, was extended.
On June 17, I had to leave town for a five-day trip. Expecting the pair to breed at any time, I solicited the assistance of two friends, Olivier Meyer and Claudia Mejia, in checking on the fish. Upon my return, Claudia told me that the pair had indeed spawned as she had observed the eggs that afternoon. Since the lights were already out in the fish room, I waited until the next morning to have a look at the aquarium. Though no eggs were seen, the behavior of one of the fish clearly indicated that larvae had been transferred to a bare flower pot. I couldn't actually see the fry, as the pot obstructed my view and I didn't want to get so close as to spook the guardian. What was clear was the identity of the single parent overseeing the young; it was the male! The female swam above the pot and would from time to time approach the larvae and the male, who did not object to her presence. Most of her time was spent patrolling around the nest and chasing away any potential intruders (including a larger male P. polleni).
A few hours later, Olivier confirmed to me that from the beginning it was the male and uniquely the male who had been looking after the spawn. As this pattern was different from practically all other known substrate-spawning cichlids, this was greatly surprising to him. During my absence, he made the following observations.
On June 17-18, premating behavior of the pair intensified. The male's displays and the pair's visits to the future spawning site became more and more frequent. The male completely emptied the contents of the flower pot (both gravel and Apogoneton bulbs), an operation that had begun prior to my departure. The spawning act itself was not observed, but must have occurred on June 19, as the eggs were first observed at the end of the afternoon on that day. Eggs were attached to the underside of the driftwood arching above the flowerpot. In order to lay her eggs, the female must have swum upside down; presumably the male did the same to fertilize them. The male was observed fanning the eggs in this position. The eggs were whitish, as opposed to red observed in the Ambomboa (Nourissat, 1993). Because of their position under the log, the eggs were not easy to see, and it was not possible to observe how they were attached to the wood; clearly however no long filament was involved.
On June 23, I resumed my observations shortly after the larvae had been deposited in the flower pot. The male was even more brilliantly colored than during courtship. As for the female, she was also quite colored up, including a purplish mask between the jaws and eyes. Over the next several days, the female's coloration was quite variable, appearing orange as in the male whenever she approached the larvae, but fading to brown upon leaving the nest site. The female often left the nest area to chase away other fishes or to look for food. This was not the case for the male who kept a constant contact with the fry. Only by dropping bloodworms over the nest could I entice the male to feed. A number of days later, I did witness infrequent forays from the nest by the male, but each lasted only a few seconds, their object always being to attack the largest P. polleni. In spite of the very short period of time involved, the male clearly signaled his intent to the female who took over the proximal guard of the nest. Upon his return, another signal relieved the female of her guardianship. The nature of this signal a stiffening of the body coupled with a slight jerk or twitch was very similar to that observed in other substrate-spawning cichlids, such as Amphilophus trimaculatus. However in the case of the lamena, the behaviors of the parental fish were reversed, reflecting the primary role of the male in looking after the fry.
About two days after hatching, I dared to approach the glass and by looking from above managed to see the larvae lying against the interior front wall of the pot. All the eggs seemed to have hatched. The fry were gathered in a compact mass which was sometimes partially scattered by the male's activities as he hung over them, frequently touching them with his mouth and fanning them with his pectoral fins. The larvae were large compared to those of other substrate-spawning species.
Their overall color was dark-brown with a large yolk sac which was totally absorbed only shortly before the free swimming stage. The number of larvae seemed to be 50-100, but eventually turned out to exceed 150. The free-swimming stage began the morning of July 1, more than eight days after hatching. Two-thirds of the fry were collected at this point to be reared in a separate aquarium. About half were captured by removing them in the flowerpot. The remainders were taken in a small hand-net. Defense of the young by the parents, especially the male, was extremely vigorous. My hand - not the net - was bitten, the resulting pain inducing a withdrawal reflex, which was hard to resist. The lamena has strong jaws with canines! Finally I had a good look at the fry in the rearing tank. They were fairly large (3-4 mm) with a robust forebody, reminiscent of a tadpole. Their overall size was more than double that of young Archocentrus septemfasciatus of the same age. The color pattern is very peculiar: a wide red-brown band circles the body and covers the abdomen. The head, except for a dark mark on the cranium, and the posterior part of the body are a light golden-beige, resulting in a bicolor pattern overall. Another peculiarity is a thin, oblique line - of a brilliant golden hue - above the eye, looking like an eyebrow.
Lamena fry are marked similarly to the young of Paretroplus kieneri bred by Nourissat in 1991. This should be considered an additional indication of the relationship between lamena and the genus Paretroplus. The behavior of the young of the two species is also suggestive of such a relationship. In both species the young do not "spread out" in a tank to forage individually, as young cichlids usually do when taken from their parents. Instead, they keep together, moving rapidly through the tank as a shoal. In the lamena, the young clung to the bottom for the first two days of observation. After this, the shoaling continued, but the school moved rapidly to the surface then back to the bottom in a twirling, frantic motion. During the first weeks they were never observed to pick food from the bottom or sides of the tank. Fortunately they ravenously consumed Artemia nauplii whenever they were presented. This behavior is assuredly an artifact of being separated prematurely from the parents (a conclusion shared by Paul Loiselle, based on aquarium-rearing of other Paretroplus species in the U.S.).
Some ten days later most of the young which had been left with the parents were removed by Claudia (I was again away). She had noticed that the number was diminishing, possibly as a result of predation by the P. polleni. This left the parents with fewer than ten fry, which quickly vanished. Without the young, the pair-bond broke down immediately, and the male resumed his aggressive posturing toward the female. I expected this phase not to last too long, as many cichlids quickly produce a spawn under such circumstances. However, this was not to be the case, and the fish waited until the end of August before breeding again. Once again the spawning act was not witnessed. The eggs were laid at the same site, but were better visible than the first time. The eggs appeared slightly elliptical and seemed to be attached at one of their extremities. The spawn was smaller than on the first occasion, and some eggs were obviously fungused. A small number of larvae resulted which were transfered to the pot by the male who (as before) cared for the brood alone. This time I decided to leave the young with the parents in order to observe parental care and the young's behavior after reaching the free-swimming stage. For their security, I removed the P. polleni and the goby, which had grown to threatening dimensions. In spite of the unavoidable turmoil which resulted from this action, the male lamena continued to guard the young faithfully. The freeswimming stage was again attained eight days after hatching. Two days later, the fry under the close guard of both parents, rose to the upper areas of the aquarium. Their behavior was totally different from that of the brood which had been separated from the parents. Although they still moved rapidly at times, the swirling, frantic motion did not occur. The young spent most of their time picking at algae growing on the upper rear glass and on the driftwood situated highest in the tank. Both parents stood guard either close by or among the fry, the female assuming nearly the same bright orange coloration as the male. The latter, however, retained greater control, as the young responded to him more frequently than to the movements of the female. During the next several weeks, the male remained in the flowerpot with the young at night. About 30 minutes before the lights were turned off (without gradual dimming), the male would begin to assemble the young and lead them into the nest. During this time, young were often taken into the mouth of the male and spat into the pot. Similar behavior was observed during the day when fry straggled away from the group with either parent performing he retrieval.
Young were never observed to feed on mucus produced by he parents as known for Symphysodon and Etroplus maculatus. Parental care lasted for nearly two months after which interest in the young declined as signs of another spawning became evident. This happened quite suddenly, resulting in a huge spawn, the eggs placed at the same site as previously. Mysteriously, the eggs quickly disappeared and no larvae resulted, the pair-bond breaking down once more. The presence of a few young from the previous brood which could not be captured and removed may have been responsible for this failure. No further spawnings have been observed, though the pair to this day is still living. In spite of their voracious appetite, growth rate of the young has been comparatively slow. Sixteen-month old fry now measure 5-8 cm. However since the age of ca. two months, signs of sexual maturity have become evident with individuals taking on an orange color and defending territories. On October 27 1994 a fish was observed caring for a spawn placed under an overturned flower pot. His partner was busily chasing away other lamena that approached the nesting site. The spawn was subsequently eaten and the pair has lost its breeding colors, but other pairs are showing signs of spawning. At present I don't have the space to isolate new pairs; the tanks now holding my F1 lamena are probably too small (and densely populated) for successful breeding to take place. I hope to remedy this situation in the near future; in the meantime I'm very relieved to learn that lamena have been bred in ponds in the U.S. Because of their unique reproductive behavior, I have felt justified in giving a detailed account of my experiences breeding the lamena. The most remarkable trait of the species is, of course, the unique paternal egg and larval care observed. To my knowledge, no other such case is known for substrate spawning cichlids. Barlow (1991) suggests that the male might be the main caretaker in Etroplus maculatus, which is interesting because members of the genus Etroplus from India and Ceylon are closely-related to Paretroplus and the lamena of Madagascar. However, contrary to the condition seen in the lamena, the female E. maculatus participates to a certain extent in the care of eggs and wrigglers. Most species of Paretroplus have now been bred in captivity, but no information on behavior is available as fish were bred in large, outdoor tanks or ponds. It will be interesting to determine whether these species show the same breeding behavior as the lamena.
If confirmed, the uniquely paternal egg and larval care of the lamena should be of great interest to scientists studying the ethology and phylogeny of the family Cichlidae. These traits will probably be interpreted as surviving within this species as a primitive, ancestral condition, supporting the hypothesis that cichlids have evolved from a marine form in which the male cared for the young (Stiassny, 1993). as such, the lamena can be regarded as a sort of missing link within the evolutionary lineage of the cichlids, meaning that the lamena and the hypothetical "proto-cichlid" most probably shared the trait of paternal care giving of the eggs and non-swimming young.
Both a beautiful and interesting species, the lamena is undoubtedly one of the most remarkable cichlid discoveries of this century's last decade. If the species can be produced commercially, it is likely to become a popular addition to the hobby. The establishment of a large and stable captive population is very much to be hoped for as the species' future in its natural habitat is uncertain. Habitat degradation and competition from introduced forms are threatening the survival of the lamena, though due to its rheophilic nature the danger is perhaps not as acute as in the case of several other endemic cichlids of Madagascar.
- Barlow, G. W.; 1991; Mating systems among cichlid fishes. In Keenleyside, M. H. A. (ed.). Cichlid Fishes: Behavior, ecology and evolution. Pp. 173-190. Chapman and Hall, London.
- Stiassny, M. L. ; 1993; Cichlids are different!; TFH (March) pp: 84-98.