The Neolamprologus brichardi complex

(This article was originally published in "Cichlids Yearbook Volume 3, Cichid press; 1993; pp. 6-13. It is here reproduced with the permission of Ad Konings, Cichlid Press.).

When the first Neolamprologus brichardi were exported from Lake Tanganyika in 1971 (Brichard, 1989) they received the apt trade name of "Princess of Burundi." These elegant cichlids were collected in Burundi near a place named Magara and although they didn't possess the bright coloration of other, at that time common, aquarium fishes, their marvelous finnage made the Princess one of the most popular cichlids among aquarists. It proved to be an excellent aquarium resident even though it is a territorial species, which defends its domain with quite some persistency. Given the right amount of room a pair will soon breed and produce successive broods of offspring. This is the most pleasant part of maintaining this species because older juveniles actively help their parents in defending younger fry (Staats, 1972). This also occurs in the lake where large aggregations of breeding adults inhabit a certain part of the rocky habitat. Most adults live and behave like monogamous pairs; it is the sheer number of pairs that sometimes gives the impression that N. brichardi is a schooling cichlid.

In 1952 the Princess of Burundi became scientifically known as a subspecies of N. savoryi, namely as Lamprologus savoryi elongatus (Trewavas & Poll, 1952). In 1974 Poll raised the Princess of Burundi from subspecies to species level. Realising that the name L. elongatus was already occupied - Lamprologus elongatus (now Lepidiolamprologus elongatus) had been described by Boulenger in 1898 - he named the Princess of Burundi Lamprologus brichardi in honour of the late Pierre Brichard who had discovered numerous Tanganyika cichlids. In 1985 Colombe and Allgayer published their "revision of the genus Lamprologus in which the Princess of Burundi was moved to Neolamprologus and Boulenger's elongatus to the revived genus Lepidiolamprologus. They argued that now that Boulenger's elongatus was no longer in the same genus as that of the Princess the old subspecific name of elongatus could be used again. For just one year the Princess was named Neolamprologus elongatus until in 1986 Poll published a revision of all Tanganyika cichlids. He accepted the new generic name Neolamprologus of Colombe & Allgayer but changed the scientific name of the Princess back to N. brichardi. His reason was that the name N. elongatus was already occupied by yet another elongatus. In 1909 Steindachner described under the name of Julidochromis elongatus a cichlid which was removed by Boulenger to the genus Lamprologus in 1915. L. elongatus had already been described so Boulenger had to rename Steindachner's elongatus. He named it Lamprologus steindachneri. Poll assumed that Steindachner's elongatus was a member of Neolamprologus which meant that that name should have priority over that of the Princess of Burundi, which therefore once again became Neolamprologus brichardi (Poll, 1986). In 1988 I found out that in all probability Steindachner's elongatus - the type of Julidochromis elongatus had reportedly been lost - was a synonym of Lamprologus callipterus, for this reason I changed the name of the Princess back to N. elongatus (Konings, 1988). Fortunately Burgess (1988) put an end to the Princess' misery by simply noting that the name elongatus was not available when it was given to her for the first time in 1952. As he pointed out, this was a mistake because the name elongatus had already been used for Boulenger's L. elongatus in 1898 and also subspecific names compete with specific names for priority. So now, once and for all, the Princess of Burundi is scientifically known as Neolamprologus brichardi.

Simultaneously with the description of L. savoryi elongatus another subspecies was described, namely L. s. pulcher. The differencebetween the two subspecies is based mainly on the pattern of the markings on the gill-cover. Both species have two dark bars between the eye and the outer edge of the gill-cover. In N. brichardi these bars have roughly the shape of the letter T lying on its side (see photograph). In N. pulcher the bar directly behind the eye is not horizontal but curves downward and does not merge with the vertical bar on the edge of the gill-cover. Brichardi-like cichlids have been exported from Zambia, the southern part of the lake. Some of these fishes were sold as Lamprologus pulcher. It is, however, not known whether these were caught at or near the type locality (which is unknown) or just looked like the holotype of L. savoryi pulcher.

Over the years Horst Walter Dieckhoff and I have gathered a good number of photographs of many different populations of N. brichardi like cichlids from all around the lake. Comparison of these pictures showed me that those gill-cover markings differ slightly between all populations. Even the population at Kasoje, Tanzania, don't show the exact markings as drawn for the holotype of L. savoryi elongatus which was reportedly collected at this locality. It thus seems that the markings may differ in the preserved state from those in the living fish.

One species in the N. brichardi complex is easily recognisable and does not show (up to now) geographical variation, i.e. Neolamprologus savoryi. This small species, with a maximum size of about 8 cm, is identified by the broad vertical bars on its body. It was found at every rocky locality visited. It prefers a depth between 10 and 40 metres which is somewhat deeper than that for N. brichardi, which is seen mostly between 5 and 25 metres. N. brichardi, and all the other races or species in the complex with markings on the gill-cover, occurs in large groups. N. savoryi, however, is usually found in pairs or solitary and is frequently seen close to groups of the other species of the complex. In the last 20 years many different N. brichardi-like cichlids have been exported under various names. We have seen Neolamprologus "Kasagera", "Daffodil", "Walteri", "Mbitae", "Black Brichardi", "White Tail" or "Palmeri", "Cygnus", and others which have not yet been mentioned in the aquaristic literature. All these variants or species are reportedly collected at different locations.

The first reports that, besides N. savoryi and N. brichardi, another species may inhabit the same biotope came from Brichard (1989). After many years of collecting at Magara, Burundi, he found that another brichardi-like cichlid lives at levels deeper than 15 metres. This species, named Neolamprologus falcicula (Brichard, 1989), lives in pairs or in very small groups, usually not far away from large groups of N. brichardi. The most irnportant difference between these species is the lack of gill-cover markings in N. falcicula. The cichlid with the trade name Neolamprologus sp. "walteri", found in Kigoma Bay and at Cape Kabogo, is a geographical race of N. falcicula. At present Neolamprologus sp. "Cygnus" from Cape Mpimbwe, Tanzania (see Cichlids Yearbook vol. 1), is also regarded as a geographical race of this species. The orange-yellow dorsal and anal fins of the juveniles of the Magara population of N. falcicula are somewhat reminiscent of the wonderful juvenile colours of the Cygnus. N. falcicula was never seen in large groups except in the case of the Walteri in Kigoma Bay, but here too they hover never more than 10 cm above the substrate, whereas N. brichardi may be seen one metre above the rocks.

In his book "Cichlids and all the other fishes of Lake Tanganyika", first published in September 1989, Brichard mentions of other localities where N. brichardi-like cichlids share the habitat with a similar species. These localities are on the southwestern shores of the lake. In his book Brichard describes an additional four new species, all from this area. Heinz Biischer, who has visited this part of Lake Tanganyika many times, also reports (1989) on a third or possibly fourth species sharing the habitat with N. savoryi and N. brichardi (and another brichardi-like cichlid). He described this species as N. marunguensis which must be regarded as a synonym of N. crassus, described several months earlier by Brichard. Brichard uses several morphometric characters to distinguish between these species, which is of course the normal practice in such descriptions. When we look at these new species we may notice that the gill-cover markings differ from species to species. Personal observations in the natural habitat indicate that the individuals of any one population do not show any apparent variation in the pattern of these markings. Therefore we can take these markings as a diagnostic feature in the identification of any of these species.

Brichard also reports that at Zongwe, Zaire, N. brichardi shares the habitat with N. splendens. Latter species - its trade name is "New Black Brichardi" - has distinct markings on the gill cover and can thus easily be recognised and distinguished from N. brichardi. But I could not find cichlids with N. brichardi-type markings on the gill-cover at Zongwe. I saw three species of the brichardi complex at Zongwe: N. savoryi, N. splendens, and N. gracilis. N. splendens is a dusky coloured cichlid. The gill-cover rnarkings are in the pattern of a V and are not much different to those found in N. savoryi. Buscher showed me a photograph of a juvenile N. splendens, which has vertical bars not unlike those of juvenile N. savoryi. N. splendens may therefore be more closely related to N. savoryi than to N. brichardi. Its maximum size is about 8 cm. Although it is common in the rocky habitat it was not seen in huge numbers as is sometimes the case with N. brichardi. The photograph on page 147 of "Tanganyika Secrets" (Konings & Dieckhoff, 1992) shows N. brichardi, and not N. splendens as indicated.

N. gracilis lacks gill-cover markings and is further recognisable by the very long filaments on the unpaired fins. Some sub-adult individuals have caudal fins almost as long as the standard length of the fish. This is the main feature that sets the species apart from N. crassus, which is much stockier and lacks the filamentous fins. This species, like N. gracilis, lacks the markings on the gill-cover. It is a rather small species with a maximum size of about 7 cm. Its type locality is at Lunangwa, but it is also found at Kapampa and in Moliro Bay, Zaire, and at Cape Chipimbi, Zambia.

The very interesting community at Kapampa, Congo, has already been discussed, with regard to N. caudopunctatus and N. leloupi, in a previous volume of the Cichlids Yearbook (1992). At Kapampa, where these two species share the habitat, N. caudopunctatus has a yellow dorsal fin, perhaps to distinguish itself from N. leloupi. A similar situation exists between N. crassus and N. gracilis. Both these species lack gill- cover markings, but N. crassus has broad white-bluish edges to the dorsal and anal fins (Biischer, 1989). The filamentous tips of the caudal fin are white in both species although they are much longer in N. gracilis. The latter species is more elongate than the more stockily built N. crassus. At Kapampa N. crassus lives at a rather deep level of the rocky biotope. Biischer (1989) reports that it was found mostly between 25 and 35 metres. N. gracilis was also observed at the same location but in shallow water at about 10 metres.

Neolamprologus sp. "White Tail" or "Palmeri" is found at Kibwesa, Tanzania (Dieckhoff, pers. comm.). This cichlid is without doubt a geographical race of N. gracilis. It is not known whether another brichardi-like species is found in the same habitat or not.

At all other known localities where three species of the complex share the habitat, there are N. savoryi, a brichardi-like cichlid with markings on the gill-cover, and one without any such markings. It thus seems that the possession of these markings or the lack of them is the main feature which segregate the species in the complex (apart from N. crassus and N. gracilis at Kapampa).

The types of Brichard's descriptions are deposited in the Museum voor Midden Afrika, Tervuren, and the four species from Zaire (N. splendens, N. gracilis, N. crassus, and N. olivaceous) all have the type locality Masanza on the label ("2nd bay north of Masanza"). In his description of N. olivaceous (also spelled olivaceus in the same publication) Brichard (1989) mentions the bay of Luhanga (=Lunangwa), Congo, as the type locality. At Kiku, which is north of the Lunangwa River, I could find only N. brichardi and N. crassus. The photograph on page 375 of his book shows a specimen of N. olivaceous which agrees with the preserved types and with the brichardi-like cichlids I found at Cap Tembwe, Kitumba, and M'toto. It is therefore possible that either the type locality lies south of the Lunangwa River (which could not be confirmed by Biischer (pers. comm.) or the designation of the type locality as Lunangwa (Luhanga) is an error.

The characteristic markings of N. olivaceous consist of two vertical curved bars on the gill-cover (like chevrons). The bar directly behind the eye borders the pre-operculum just as the second bar borders the outer edge of the gill-cover. This pattern is characteristic of N. pulcher. Although the type locality of N. pulcher is unknown the gill-cover markings of this cichlid are seen in several populations. I therefore propose to regard N. olivaceous as a synonym of N. pulcher.

N. pulcher is a small cichlid (maximum size about 6 cm) which lives in small groups in the somewhat deeper rocky habitat. Most individuals were seen below 10 metres. Juveniles at Cap Tembwe, Zaire, have orange dorsal fins. Neolamprologus sp. "Daffodil" found at Kalambo, Tanzania/Zambia have similar gill-cover markings and yellow unpaired fins in juveniles as well as in adults. Because of the apparent importance of the pattern of the gill-cover markings I consider the Daffodil a geographical race of N. pulcher.

N. brichardi seems to have the widest distribution of all the species in the complex. I regard all brichardi-like cichlids with T-shaped markings on then gill-cover as geographical variants of N. brichardi. V-shaped markings are found in N. splendens and chevron-shaped markings in N. pulcher. N. falcicula is recognisable by the lack of markings on the gill-cover and by the black outer edges of the dorsal and anal fins. N. gracilis lacks the markings as well but has white edges to the unpaired fins. It differs from N. crassus in its elongated body, the depth of which is 25 to 28% of the standard length (Brichard, 1989). The comparable percentage in N. crassus is 30 to 35%.

It is interesting to note that thus far N. brichardi, N. gracilis, and N. pulcher have a discontinuous distribution and are found on the west as well as on the East Coast of the lake.

References

• Brichard, , 1989, Cichlids and all the other fishes of Lake Tanganyika. TFH, Neptune, U.S.
• Buscher, , 1989, Ein neuer Tanganjika-Cichlide aus Zaire. Neolamprologus marunguensis n. sp. (Cichlidae, Lamprologini). DATZ 42, pp: 739-743.
• Burgess, W., 1988, The many faces of Lamprologus elongatus. TFH, Nov. pp: 61-62.
• Konings, , 1988, Lamprologus callipterus en Julidochromis elongatus. NVC periodiek, (Dutch Cichl. Assn.) 14 (2).
• Konings, A. & Dieckhoff, H.W., 1992, Tanganyika secrets. Cichlid Press, St. Leon-Rot, Germany.
• Staats, , 1972, Lamprologus savoryi elongatus, Trewavas & Poll 1952 (Prinzessin von Burundi). DCG-Info (German Cichl. Assn.) Nr. 10; pp: 85-90.
• Trewavas, E. & Poll, , 1952, Three new species and two new subspecies of the genus Lamprologus , cichlid fishes of Lake Tanganyika. Bull. Inst. Sc. nat. Belgique. Vol. 28 (50); pp: 1- 16.