The Volume 2

CICHLIDS yearbook

Ad Konings (Ed.)

100
4
 THE
CICHLIDS
YEARBOOK
Volume 2

Ad Konings (Editor)

1
 Cover photographs:
1 - Protomelas sp. “Steveni taiwan”, Lake Malawi, Malawi.
1 2 - Tilapia tholloni, Alima River, Congo.
3 - “Cichlasoma” sp. “Labridens Tamasopo”, Pánuco, Mexico.
4 - Tropheus moorii (Murago), Lusingo, Lake Tanganyika, Zaïre.
2 3 4

Text and photographs by Ad Konings

except as otherwise indicated

Mary Bailey (Crediton, UK)

corrected the manuscript

The editor wants to thank the following persons who

supplied various cichlids for photographic purposes:
Peter Baasch (Stegen, Germany)
Marc Danhieux (Maltavi, Hohenahr-Erda, Germany)
Alain Gilot (Zaïre Cichlids, Kalemie, Zaïre)
Stuart Grant (Salima, Malawi)
René Krüter (Krüter Tropicals, Rotterdam, Netherlands)
Roland Numrich (Mimbon Aquarium, Köln, Germany)
Edwin Reitz (Aquapport, Ronnenberg, Germany)
Dirk Verduijn (Verduyn Cichlids, Zevenhuizen, Netherlands)

Distributors:

USA: Old World Exotic Fish, Inc., P.O.Box 970583, Miami, Florida 33197
UK: Animal House (U. K.), Ltd., QBM Business Park, Birstall, Batley, West Yorkshire WF17 9QD
Sweden: Fohrman Aquaristik AB, Odds Väg 7, 433 75 Partille
Germany: Aquapport (Edwin Reitz), Köselstraße 20, 3003 Ronnenberg
Netherlands: NVC, Lieshoutseweg 31, 5708 CW Stiphout

ISBN 3-928457-05-5

Copyright © 1992. Cichlid Press. All rights reserved.

No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means—
electronic, mechanical, photocopying, recording or otherwise—without the prior permission of the authors, and the pub-
lisher.

Cichlid Press, Blütenweg 17, 6837 St. Leon-Rot, Germany

Printed by RAKET B.V., Pijnacker, Holland

2
 The CICHLIDS yearbook
CONTENTS
Introduction ........................................................................................................................................................... 5

Tanganyikan Cichlids
Clues to a step-wise speciation by Ad Konings .................................................................................................... 6
Spawning Xenotilapia spiloptera by Ad Konings .............................................................................................. 10
The genus Tropheus by Ad Konings ................................................................................................................... 13
Simochromis marginatus........................................ 16 Neolamprologus sp. “Ubwari Buescheri” ............. 22
Simochromis babaulti ............................................ 17 Neolamprologus sp. “Kavalla” .............................. 23
Xenotilapia ornatipinnis ........................................ 18 Cyphotilapia frontosa ............................................ 24
Ctenochromis benthicola ....................................... 19 Cyprichromis sp. “Leptosoma Jumbo”.................. 26
Neolamprologus petricola ..................................... 20 Paracyprichromis brieni ........................................ 29
Neolamprologus leloupi ......................................... 21

Malawian Cichlids
The genus Cynotilapia Regan, 1922 by Andreas Spreinat ................................................................................ 30
The Protomelas taeniolatus-complex by Ad Konings ........................................................................................ 34
Tramitichromis lituris by Ad Konings ................................................................................................................38
Stigmatochromis sp. “Modestus Makokola” ......... 41 Taeniochromis holotaenia ...................................... 48
Dimidiochromis strigatus ...................................... 42 Lethrinops micrentodon ......................................... 50
Copadichromis boadzulu ....................................... 43 Labeotropheus fuelleborni ..................................... 51
Buccochromis rhoadesii ......................................... 44 Pseudotropheus saulosi ......................................... 52
Corematodus taeniatus .......................................... 45 Labidochromis sp. “Hongi” ................................... 53
Aulonocara rostratum ............................................ 46

Victorian Cichlids
Introduction to taxonomy and ecology: Part I by Ole Seehausen ...................................................................... 54

West African Cichlids

Lamprologus sp. “Kinganga” by Frank Warzel ................................................................................................. 60
Tilapia tholloni (Sauvage, 1884) by Jan ‘t Hooft ............................................................................................... 62
Ctenochromis polli / Thoracochromis demeusii by Martin Geerts ...................................................................64

Central American Cichlids

The “Cichlasoma” labridens-complex by Juan Miguel Artigas Azas ...............................................................65
“Cichlasoma” minckleyi Kornfield & Taylor, 1983 by Ad Konings .................................................................. 71
“Cichlasoma” septemfasciatum ............................. 74 Thorichthys pasionis .............................................. 76
Theraps coeruleus .................................................. 75

South American Cichlids

Crenicichla species from the Rio Xingú by Frank Warzel ................................................................................ 77
Crenicichla sp. cf. regani ....................................... 82 Guianacara sp. “Red Cheek” ................................ 84
Acaronia vultuosa .................................................. 83 Krobia species ........................................................ 85

Cichlid Maintenance
The “Aufwuchs-feeder” by Roger Häggström ...................................................................................................86
Breeding Tropheus the natural way by Gerard Tijsseling .................................................................................. 87

Cichlid Literatim
The cichlid bible by Martin Geerts ....................................................................................................................88
Cichlid classics by Mary Bailey ......................................................................................................................... 90
Spawning techniques in mouthbrooders by Ethelwynn Trewavas & Ad Konings ............................................. 93

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4
 The CICHLIDS yearbook

Introduction

No less than eighteen authors have put together the Last but not at all the least in the long row of “new”
second volume of the cichlids yearbook. It was pleas- authors, Mary Bailey (Crediton, UK) explains scien-
ing that almost all contributors to the first volume were tists (should) give names to fish. Mary, who has been
able to write down another piece of their knowledge on the committee of the British Cichlid Association
and experience in this one. In addition eight new au- since 1982 and has studied English and Latin, was
thors have written for this volume. Another pleasing also of tremendous help in correcting the manuscripts
fact is that Horst Walter Dieckhoff (Lake Fish, Herten, of the yearbooks. She has kept and bred many species
Germany), the famous underwater photographer, is and is an aquaristic consultant for several aquarium
back in “cichlid-business”. For this volume he has pro- magazines in the UK.
vided some of his photographs but we have made some A very important publication has been issued re-
plans for the near future.... cently, namely Cichlid Fishes; Behaviour, Ecology and
The “new” authors are briefly introduced in the Evolution. It is the cichlid bible for the years to come
same order as their articles appear: for every aquarist interested in cichlids. It is discussed
Hans-Joachim Herrmann (Hamburg, Germany), a by Martin Geerts.
Tanganyika specialist and known for his book on these
cichlids, describes two new variants of Simochromis, In the previous volume I have reported on
which he has collected himself. Neolamprologus leloupi and said that N. caudo-
Another book-author and Malawi specialist, Dr. punctatus should be regarded as a synonym of this
Andreas Spreinat (Göttingen, Germany), deals with species. However, when I visited the locality in Zaïre
the genus Cynotilapia and relates his experience with where the yellow-dorsalled cichlid is collected, it was
some other cichlids as well. immediately clear that I was wrong: both species, N.
Edwin Reitz (Aquapport, Ronnenberg, Germany), caudopunctatus and N. leloupi , live sympatrically in
a professional and very experienced cichlid breeder, the same habitat. I further explain the situation on page
describes the breeding procedure of Aulonocara 21.
rostratum. With regard to Willem Heijns’ contribution in vol-
Recently the first commercial shipments of fish from ume 1 about “Cichlasoma” spinosissimum , Jaap-Jan
Lake Nyasa, as Lake Malawi is known in Tanzania, de Greef (Parrish, Florida), who rediscovered this spe-
have been exported from that country. Two independ- cies for the hobby, wrote me that he found the species
ent operations have been started. In the next volume in pools at the southern end of Lago de Izabal but not
we will have some more species for you but this time in the lake itself. Jaap-Jan collects fish in Central
Peter Knabe (Schlangen, Germany), who is a scien- America and Africa and breeds them for a hobby.
tist working in Tanzania in the Haplochromis Ecol-
ogy Survey Team and thus has first-hand experience, Without the hospitality and cooperation of several
has contributed his first instalment. exporters and friends it would not have been possible
Jan ‘t Hooft (Hendrik Ido Ambacht, Netherlands), to show you the cichlids in their natural habitat and
a lifetime aquarist and the founder of the Dutch cichlid give information on how they live. I therefore grate-
association, tells us about one of his passions, namely fully acknowledge Stuart M. Grant (Salima, Malawi),
Tilapia. Alain Gillot (Zaïre Cichlids, Kalemie, Zaïre), Gary
Do you keep mbuna with larger Malawian haplo- Kratochvil (San Antonio, Texas), Juan Miguel Artigas
chromines in your tank? Then you know that at feed- Azas (San Luis Potosí, Mexico) and Mireille Schreyen
ing time the mbuna devour the major part of the food. (Fishes of Burundi, Burundi).
It usually takes them ten seconds to do so. Roger
Häggström (Örnsköldsvik, Sweden), the editor of Ad Konings
Ciklid Bladet, the periodical of the Swedish cichlid
association, tells us how we can keep the mbuna busy
eating.

5
 TANGANYIKAN
CICHLIDS
Clues to a step-wise speciation
Ad Konings

This race of Ophthalmotilapia ventralis at M’Toto, Zaïre closely resembles that from Cape Mpimbwe, Tanzania (see page 7).

Among the many endemic species of cichlids in Lake substrate-spawners like Neolamprologus caudo-
Tanganyika which are known to date, there are a punctatus and N. leloupi (for the latter species see page
number of species which have evolved geographical 21).
races. The best known examples of such cichlids are Recently, Martin Geerts and I made a short expedi-
the species of the Tropheus-complex. Many different tion along the southern Zaïrean coast from Kalemie
geographical variants have been found which all seem to Moliro. It was our intention to make a preliminary
to belong to five, maybe six, species. Besides Tropheus inventory of the rock-dwelling cichlids in this part of
there are other species like Cyathopharynx furcifer and the lake. During our trip we became aware of a re-
Ophthalmotilapia ventralis which have evolved sev- markable feature which may have a greater impact on
eral different races. Not only the mouthbrooders among the theory of speciation than our first observations
the cichlids have formed distinct populations but also indicated. We observed geographically isolated

6
populations of species whose coloration had a remark- part of the Zaïrean coast (see photo page 8 top) is prac-
able resemblance to populations along the opposite tically identical to the one on the other side of the lake
side of the lake. We knew about these populations (see photo page 8 bottom). In the Zambian part, how-
through the many photographs made by Walter ever, another race inhabits the intermediate and sandy
Dieckhoff in the Tanzanian part of the lake. biotopes (see photo page 89).
Ophthalmotilapia ventralis inhabits the southern The geographical variant of T. moorii found between
half of the lake and is known in about a dozen distin- Moba and Zongwe closely resembles the small popu-
guishable colour varieties, each geographically iso- lation at Kibwesa, Tanzania (see Konings, 1990 TFH
lated. On the Tanzanian coast at Cape Mpimbwe lives 39 (3): 71) while the race found at Kapampa (see photo
a race which is characterized by a black-blue ground page 9) shows a strong resemblance to the race near
colour with an irregular, broad white band running di- Mpulungu. The latter two populations, however, are
agonally across the body (photo this page). separated by a long shoreline which is inhabited by
Between Cape Tembwe and the bay of Zongwe on about a dozen different other variants.

Ophthalmotilapia ventralis at Cape Mpimbwe, Tanzania.

the Zaïrean coast an almost identical variant of O. ven- T. polli has been described from the central Tanza-
tralis is found (see photo page 6). Further south along nian coast and is characterized by having four spines
the coast a yellow race of O. ventralis inhabits the in the anal fin, a feature it shares, among its conge-
shallow rocky biotope. The race found at Kapampa ners, only with T. annectens. Other Tropheus have 5
closely resembles that found at Kipili, Tanzania to 7 spines. At M ’Toto, Zaïre, I observed T. annectens
(Konings, 1988: 61). At Lupota, Zaïre, the variant in its habitat and found no apparent difference in be-
shows more yellow pigment and is almost identical haviour or morphology to that of T. polli.
with that found at Malasa Island, Tanzania. Neolamprologus leloupi occurs on both sides of the
Cyathopharynx furcifer is known from most loca- lake (in the southern half) but is not found in Zambia.
tions around the lake and about six different ’ races On page 21 some of its affinities with N. caudo-
are known. The race found along most of the southern punctatus are discussed. Both at Kapampa, Zaïre (see

7
Cyathopharynx furcifer,a male photographed south of Moba, Zaïre.

Cyathopharynx furcifer at Kipili, Tanzania.

8
photo page 21) and at Kalambo, Tanzania/Zambia
(René Krüter, pers. comm.) an identical race of N.
caudopunctatus – which is not known to occur any-
where else — lives sympatrically with N. leloupi at
the borderline of both species’ distribution.
There are several other species which have similar
(not identical as in the previous examples) popula-
tions on both sides of the lake, e.g. N. sexfasciatus,
several species of the N. brichardi-complex and
Chalinochromis sp. “Popelini” in Zaïre and C. sp.
“Bifrenatus” in Tanzania. With regard to the follow-
ing discussion, it is important to realize that different Tropheus moorii at Kapampa, Zaïre.
races of the species mentioned inhabit the Zambian
waters. The first question one asks is how come such
isolated populations look so much alike when it is all! These rather basic observations may lead us to
known that they can develop many other colour pat- conclude that when the bulk of the individuals of a
terns as well. The east and west coasts in the southern population remain together in one large breeding
part of the lake are separated by the deepest water group, the cichlids of that group will not change, prob-
found in Africa. If one is conversant with the current ably not in a million years! But when a drastic change
theories about the fluctuations in lake levels since the does occur and new populations are founded, a new
origin of the lake, one can easily imagine a period in species could be developed almost overnight.
which the water dropped to a level which was low One could envisage the development of a new popu-
enough to connect parts of opposite coasts. In such lation as follows. As the rising water level opens up a
small paleo-lakes the aforementioned species could new, suitable, habitat, wanderers from a nearby popu-
have been present as a single geographical popula- lation could cross uninhabitable areas and settle at the
tion. The rising water caused the fish to move upwards new location (horizontal migration). Of course not just
in order to keep up with the suitable habitat (vertical one species would occur at such a new site but many
migration). This eventually resulted in the populations different species would found a new population. When
on either side becoming isolated. This seems to me such new populations are founded by small numbers
the only plausible way to explain apparently identical of individuals then there is a statistical chance that a
races at both sides of the lake. new colour variant will develop (interbreeding). Im-
If this is indeed the only reasonable explanation then portant factors in the development of a new race are,
its implications are very important. During the low firstly, the number of wanderers founding the new
water stand there was no water in the Zambian area. population. Secondly, the constitution of the local
Therefore all Zambian populations must have devel- fauna and the local circumstances may greatly influ-
oped from individuals from the more northerly regions ence the success of a particular species. The latter two
(horizontal migration). The fact that they almost all factors probably act as catalysts in speciation and the
have a different colour pattern leads to the conclusion development of a new variant may depend solely on
that these races must have developed in Zambian wa- the number of individuals at the site.
ters. We may then further conclude that speciation When a balance between the available species,
takes place mainly during the initial development of a number of individuals, and available niches has been
new population. This, however, is at variance with established “evolution” is suspended until a new ma-
Darwin’s theory that evolution is a continuous proc- jor change occurs in this habitat.
ess whereby new species are slowly generated.
Combining the existence of the similar variants on Acknowledgement
both sides of the lake with the widely accepted theory
of the fluctuations in lake level, one can only envis- I was greatly stimulated by the discussions I en-
age a step-wise evolution in the case of Tanganyikan joyed with Martin Geerts and Mary Bailey. I sincerely
cichlids. This hypothesis is further supported by the thank Alain Gillot of Zaïre Cichlids for giving us the
“fact” that during the development of the new popu- opportunity to make a survey along the Zaïrean coast.
lations in the southern part of the lake (while the wa-
ter level rose) the old populations did not change at

9
 Spawning Xenotilapia spiloptera Poll & Stewart, 1975
Ad Konings

The genus Xenotilapia can be divided into two groups; ing the substrate for something edible. It may inter-
one group consists of maternal mouthbrooders and the mingle with other species to form larger schools but
other of species which form pairs during the breeding most often groups consisting only of X. spiloptera are
period and which employ the biparental mouth- observed.
brooding technique. However, this subdivision may With the approach of the breeding season – at
not always be as clear-cut as is stated here (see page present it is not known whether there is a regular an-
18, Mark Smith’s article about X. omatipinnis). nual breeding season, or whether breeding is triggered
X. spiloptera belongs to the group of biparental mouth- by the effect of some external stimulus on the mem-
brooders. The holotype of this species was collected bers of the school – the school moves closer to the
at Nkumbula Island near Mpulungu in Zambia, but its rocky habitat and splits up into pairs. Each pair stays
distribution seems to spread along other parts of Zam- in a territory which is defended mainly against
bia and along the Zaïrean and Tanzanian shores. conspecifics.
During the breeding period X. spiloptera is restricted The pair bond is established by repeated courtship

A mouthbrooding female Xenotilapia spiloptera from Kipili, Tanzania.

to the rocky habitat. This has led to the existence of displays by the male as well as by the female. In the
several geographical races. Most races have a colorless artificial environment of the aquarium it seems that
dorsal fin with some black markings on its edge. The the bond between the pair is strengthened by the con-
race at Kigoma has a dorsal fin with tiny colored spots tinuous protection of the territory. If only one pair is
but lacking the blotchy markings found in all other kept in the tank there is a risk that the male and female
known populations. The races inhabiting the rocky will quarrel and that one of them, not always the fe-
shores near Kipili in Tanzania have an attractive yel- male, might end up cowering in one of the tank’s cor-
low coloration in addition to the black markings on ners. Even when the pair is not brooding, male and
the dorsal fin. female stay together in a relatively small area.
When not breeding, X. spiloptera sometimes lives The four pairs I keep in an 800 litre aquarium are
in large schools over the sand. Here it forages by sift- each satisfied with an area about 30 cm in diameter.

10
The female has just laid some eggs; the male waits behind her.

When one pair decides to spawn, it defends an area The female will not lay any eggs if the male is not
about double that size. It is difficult to predict an im- behind her. While he is chasing intruders from the ter-
minent spawning but mutual courtship increases no- ritory she may remain at the site but she may also join
ticeably, sometimes days before the actual spawning. the male in the territorial defence. After a short inter-
A slight change in the color pattern occurs as well; ruption the female again lowers her body to the
female as well as male acquires black markings in the substrate and waits for the male to take position be-
upper and lower part of the iris. These, together with hind her before she lays a new batch of eggs.
the pigment in the eye, form a black vertical bar across Spawns may be as large as 40 eggs but much seems
the eyeball. I don’t know if this is a sign of readiness to depend on the buccal capacity of the female. If the
to spawn, but it is seen mostly when the pair is en- female’s mouth is getting full she has to shuffle the
gaged in spawning or shortly before the act. eggs around before she can pick up new ones. Usu-
Spawning takes place inside the pair’s territory but
there is no specific site or nest where the eggs are de-
posited. In fact, the deposition site may change dur-
ing spawning. In the maternal mouthbrooder group of
Xenotilapia, male and female circle around each other
before the eggs are laid, but X. spiloptera starts spawn-
ing when the female suddenly deposits some eggs on
the substrate. I have never noticed a signal from the
male for the female to start. He usually waits behind
her, about 3 cm above the substrate, until she clears
the site and leaves the eggs to be fertilized by him.
While the male positions his vent over the eggs to fer-
tilize them, the female turns around and waits until
the male moves away. Then she picks up the eggs. The male fertilizes the eggs when they are on the substrate.

11
ally the male has his head near the eggs when the fe- four weeks taking care of one brood, males of a ma-
male collects them and sometimes picks up some eggs ternal mouthbrooder can fertilize eggs of many females
himself. Once I noticed that he picked up two eggs, in the same period. In theory, this should not make a
which the female apparently didn’t collect quickly difference as long as there are as many females as
enough, and swam away while chewing on them. Then males. The drawback of biparental mouthbrooding is
the pair went through another cycle, and when the fe- the fact that a territory has to be maintained by the
male wanted to pick up the new batch of eggs the male pair. In maternal mouthbrooders only the strongest
spat the two eggs in front of the female’s mouth. They males occupy a territory, and the mouthbrooding fe-
were immediately picked up by the female. During males gather in nursery schools. Where population
the next round, the female’s mouth was probably too density is high, as is the case in Lake Tanganyika, this
full and it took so long before she had arranged the offers a distinct advantage. Instead of having many
eggs already inside her mouth that the male again took territories spread over a large area, a species can propa-
some eggs, but this time he ate them. The female didn’t gate in a relatively small region of the biotope. Only
produce more eggs after this incident. the males have to worry about a territory and since
Shortly after spawning the pair reduces the terri- there are fewer needed a loss during the selection proc-
tory to the usual 30 cm, but this is defended with more ess probably enhances the viability of the species.
energy than usual. The first nine to twelve days the The stimulus for the female of the biparental
female broods the embryos and refrains from feeding. mouthbrooder to transfer the larvae to the male’s
After this period the larvae are transferred to the male’s mouth may be hunger. During the 9 to 12 days incu-
mouth. I have never witnessed the procedure in its bation she does not eat at all. Females of a maternal
entirety but it is likely that, after a set ritual, the fe- mouthbrooder endure a longer period without food but
male spits all the larvae in front of the male’s mouth. may eat small morsels in the second half of the incu-
The male retrieves them quickly and continues the bation period. Such mouthbrooding females also
brooding for another ten days. exspend much less energy than those of a biparental
When the brooding is completed the male releases mouthbrooder which also have to defend territory.
the fry in the territory where they are protected by In Lake Tanganyika a maternal mouthbrooder is
both parents. For the first few days the fry may take better off in several respects. This could explain the
refuge inside the male’s mouth but most of the time observation that breeding X. spiloptera are found at
they sit on the substrate. They sometimes wander off deeper levels of the biotope than are maternally
into other territories where they are protected by the mouthbrooding Xenotilapia. Breeding pairs are usu-
resident pair. The fry measure about 15 mm at the time ally found at depths of between 15 and 40 meters .
of release and it takes them about two years before
they have reached the adult size of approximately
10 cm.
The biparental mouthbrooding technique is not more
advanced than the maternal mouthbrooding procedure.
While mouthbrooding pairs have to devote three to

After fertilization the eggs are collected by the female.

12 The male watches the female picking up the eggs.

 The genus Tropheus
Ad Konings

The “Murago Moorii” exists in several variants. Here a male of the “Yellow Murago” collected near Zongwe, Zaïre

The species of the genus Tropheus have for many years ing unicellular diatoms from the algae strands, par-
been some of the most popular cichlids among tially clear the biocover from sediment and make it
aquarists. There are many hobbyists who keep exclu- thus more suitable for Tropheus to feed on (Takamura,
sively Tropheus species. Although there are only six 1984).
known species, at least four of these show great geo- In the artificial environment of the aquarium T.
graphical variability. Fortunately most aquarists are moorii, from the southern part of the lake, and T. sp.
aware of this fact and rarely house different variants “Black”, form the northern, behave as strongly terri-
of the same species in one tank. This would lead in- torial cichlids. Typically the largest males will divide
evitably to bastardization. the available space in the tank among themselves and
The members of the genus Tropheus are usually well leave the border areas of their territories as living space
represented on most rocky coasts. Except for T. for females and weaker males. Territories in the
duboisi, they all inhabit the upper 10 metres of the aquarium measure more than 60 cm in diameter. In
biotope. Not only for their protection but also for their nature, the territoriality of these two species is not al-
food, Tropheus species are restricted to areas with ways as evident as it is in a tank. Although males have
rocks. The feeding habits of T. duboisi have not been a territory where spawning takes place and in which
extensively investigated, but those of the other spe- they forage, conspecifics are not always chased from
cies have. T. moorii, T. brichardi, T. polli and T. sp. the premises. Sometimes we may observe a group of
“Black” are all grazers which shear the filamentous about 20 (T. sp. “Black”) to 100 (T. moorii) individu-
algae from the rocky substrate (Yamaoka, 1983). T. als grazing on the same rock. In the wild, T. brichardi
annectens, which is restricted to the western, Zaïrean, and T. polli are permanently territorial and, especially
shores of the lake, is closely related to and maybe even T. brichanti, are difficult to keep in small numbers in
conspecific with T. polli, and behaves similarly to that an aquarium.
species. It seems that Tropheus has a feeding relation- Many aquarists have great interest in maintaining a
ship with several species of the genus Petrochromis. breeding group of a particular variant of one of the
Members of the latter group, which are browsers comb- species. Not only for the continuous interactions be-

13
tween the members of the group but also to spread the maintaining Tropheus.
aggression of the most dominant males, it is advis- Wild caught fishes and fishes from aquaria in which
able to have as large a group as possible. If each indi- wild caught specimens are housed have many differ-
vidual is given 25 to 40 litres of water a balanced breed- ent kinds of parasites. In the wild, these parasites live
ing group can be established in a suitably decorated in balance with the host’s immune system and are
aquarium. Caves are not necessary as they will not rarely present in harmful quantities. By virtue of the
provide shelter for harassed individuals; only the domi- presence of a low number of parasites the host con-
nant males will occupy caves. Stones should be placed tinuously maintains its resistance against an explosive
in such a way that clearly separate groups are created. increase in numbers of that parasite, although it never
Each group or heap of stones, if approximately 50 cm becomes totally immune. Fish in a healthy aquarium
in diameter, will be regarded as a territory by a male have very few parasites and thus mostly lack this semi-
and respected by the others. immunization. When a wild caught fish is introduced
Often more females are kept than males. This may to an established group, it is immediately weakened
work for most species of Tropheus but not always for by the repeated physical assaults it endures from all
T. brichanti. The latter species is best kept with as members of the group. This has a weakening effect on
many males as females. its immunity to parasites and a serious infestation is
Before a group of Tropheus is introduced into the liable to result. Not only may the newly introduced
aquarium all of the fish must be in good condition or fish die but the disease usually infects the nonresist-
at least be in the same condition, and all of the fish ant inhabitants as well. A single fish may thus com-
must be introduced at the same time. The introduction pletely wipe out a long established breeding colony.
of additional specimens to the group at a later stage Therefore, never introduce untreated wild caught fish
must be avoided at all times! Newly introduced fish into a group of Tropheus.
more often than not upset the established hierarchy in Several parasites are specific to a group of cich-
the group and frequently end up being chased by all lids; they affect only those species while other spe-
other individuals. If the additional specimens are wild cies in the same aquarium do not show any sign of
caught fishes which have not been quarantined long disease. Species that carry some of the Tropheus-af-
enough, an early death is generally the result. Cur- fecting parasites are Goby Cichlids (Eretmodus,
rently this is the most important problem aquarists have Spathodus and Tanganicodus), Petrochromis,

A male Tropheus brichardi from the Kavalla Islands, Zaïre.

14
Simochromis, and Pseudosimochromis. There are sev- pathogens. The faster the water is recycled through
eral ways of “cleaning” wild caught fish, but all of the trickle filter the more effectively it frees it from
them involve a quarantine period. A wild caught fish, parasites. When the volume of water passed through
living in balance with its parasite burden, can be the filter in one hour equals that of the tank’s volume,
cleansed of them by killing the existing parasite or by an optimum treatment has been achieved. It is not
preventing re-infection where the parasites have left known if faster cycles have a better or even the same
the fish in the course of their life cycle. To do this, effect. Slower cycles don’t work that well.
however, one needs to know what organisms are in- So, wild caught fish could be cured by placing them
volved. In Tropheus broad spectrum antibiotics and in a quarantine tank which is filtered by a trickle fil-
anti-parasitic solutions have had little effect in dam- ter; even when the tank is a part of a central filter sys-
ming the spread of disease. What kills most Tropheus tem! It is important to leave the fish for at least two
seems to be highly infectious agents, either virus or months on their own in that quarantine tank and have
parasites. Many aquarists have tried numerous medi- a separate hand-net for that tank only. When direct
cines to cure their fish, some with success. Reasoning water-to-water contact with other tanks is avoided the
tells us, however, that there may be several diseases result is healthy fish.
involved and that thus one medicine will not auto-
matically cure all fish. References
As mentioned before, a fish may also be cured by
preventing the parasite from re-infecting the host. If a TAKAMURA, K. (1984) Interspecific relationships of
parasite cannot find a host it will die. The infectious aufwuchs-eating fishes in Lake Tanganyika. Env.
phase may live a long time but the agent infecting Tro- Biol. Fish. 10 (4); pp 225-241.
pheus seems to have a relatively short lifespan. My YAMAOKA, K. (1983) Feeding behaviour and dental
personal experience and that of Hans Herrmann (pers. morphology of algae scraping cichlids (Pisces:
comm.) is that a biological trickle filter strongly in- Teleostei) in Lake Tanganyika. Afr. Study Monogr.
hibits the spreading of the disease. This can be ex- 4; pp 77-89.
plained by the fact that infectious parasites are carried
away from the fish into the filtration system. It is our
experience that water coming from the filter is free of

Tropheus sp. “Black”, a geographical variant from Kiriza, Ubwari Peninsula, Zaïre.

15
 Simochromis marginatus Poll, 1956
Hans-Joachim Herrmann

A wildcaught male Simochromis marginatus from Nyanza, Burundi. Photo by Hans-Joachim Herrmann.

Since its description 35 years ago, this cichlid has been pared them with the original descriptions of the two
known to the scientific world but has never been in- species could I conclude that I had collected S.
troduced into the aquaristic hobby. Its rather unexciting marginatus. The only difference is the relatively deeper
coloration could be one of the factors that postponed caudal peduncle of S. margaretae.
its entry into the hobby; on the other hand the first Like all other species of the genus S. marginatus
few specimens were only recently collected alive by spawns easily in captivity. The male constructs a shal-
some German aquarists. Until then its only known low spawning-dish in the sand and leads ripe females
occurrence was located on the northwestern coast of with undulating and vibrating movements into his nest.
the lake, along the Zaïrean coastline. Now a second When both partners are ready to spawn they circle the
location has been found on the Burundi coast in the spawning-site. The eggs are fertilized inside the fe-
northeastern part, just north of Nyanza. male’s mouth. When all eggs are laid the female re-
The fact that I discovered Simochromis marginatus treats from the male’s territory and cares for the eggs
in the rocky habitat at this location made me a little during the following three weeks. After this period all
uneasy. This part of the lake, which is easily accessi- egg yolk has been absorbed and the fry are about 1 cm
ble, has been thoroughly investigated by the late Pierre long. Only when danger threatens are they taken back
Brichard. This author has denied, in his book “Cich- inside the female’s mouth.
lids and all the other fishes of Lake Tanganyika” At a size of about 3 cm juvenile S. marginatus al-
(1989), the existence of S. marginatus in this part of ready show intraspecific aggression, and this is known
the lake. My unease is increased by the fact that S. to occur among the other species of the genus as well.
margaretae – this species has not yet been exported Such aggression can easily be reduced by accompa-
alive – is recorded from a location not far from the nying them with other non-related species like Troph-
place where I found S. marginatus, i.e. Kigoma Bay. eus moorii or a species of the genus Petrochromis.
S. margaretae is also characterized by a black mar- Males can reach a maximum size of approximately 11
ginal band in the dorsal fin and has a very similar shape cm while females, which do not show a distinct black
to S. marginatus. marginal band in the dorsal fin, remain about 2 cm
Not until I had examined the specimens and com- smaller.

16
 Simochromis babaulti Pellegrin, 1927
Hans-Joachim Herrmann

Simochromis babaulti; a submissive male is visible in the background. Photos by Hans-Joachim Herrmann.

The genus Simochromis consists of small to medium

sized mouthbrooding cichlids which inhabit the rocky
shallows or the intermediate biotopes in Lake Tan-
ganyika.
One of the smaller species is Simochromis babaulti
which has a maximum length of about 10 cm and
which is fairly well known among aquarists. Because
of its rather aggressive — intraspecific — behaviour,
and its lack of a sparkling coloration, S. babaulti is
maintained by only a few specialists.
In 1989 I had the opportunity to bring back alive a A female Simochromis babaulti from Nyanza, Burundi.
new variant of this species which had not been known
before. Males of this variant show yellow instead of
beige vertical bars. Moreover there are a number of
bright red spots sprinkled over the upper part of the
body. The latter feature reminds us of S. pleurospilus
(see photo) and possibly these two species have a much
closer relationship than has been thought previously.
Females of this variant, which have a maximum size
of approximately 8 cm, have a very attractive colour
pattern consisting of a large red blotch covering the
head, ventral fins and part of the body. The intensity
of the red pigment varies according the emotional state
of the fish. I found this variant in a shallow intermedi-
ate habitat a few miles north of Nyanza, Burundi. An adult male Simochromis pleurospilus from Zambia.

17
 Xenotilapia ornatipinnis Boulenger, 1901
Mark Smith

A courting male Xenotilapia ornatipinnis. Photos by Mark Smith

Xenotilapia ornatipinnis has recently been imported Its large, flattened eyes are also marvelous adaptations
under the name of “Pearly Xenotilapia”. The fishes for living in poorly lit environments. The maximum
currently in the hobby are collected from the northern recorded size is 13 cm total length.
part of the lake and were exported by Fishes of In the aquarium, X. omatipinnis readily adapts to
Burundi. the regular aquarium fish food. It behaves very peace-
X. ornatipinnis is known to occur down to depths of fully; never did I observe any torn fins or any indi-
150 metres (500 feet). However, it also occurs in much viduals cowering in the corners of the aquarium.
shallower waters, a fact which has enabled its collec- X. omatipinnis has been bred in the aquarium and
tion. As with several species of Tanganyikan cichlids, in most instances the female broods her embryos to
X. omatipinnis seems to display a wide depth range full term on her own. Ron Sousy (pers. comm.) found,
possibly related to the daily vertical migration of plank- however, that the female would sometimes give the
ton on which it feeds. X. omatipinnis displays a several days old embryos to the male who would then
“pearlescent” body coloration so very characteristic incubate the developing embryos to full term. On one
of deep living, sand-dwelling cichlids from the lake. occasion a carrying male gave the developing embryos
to yet another male! This foster father held for a little
more than one day before disposing of the brood. To
know whether or not this is a naturally occurring event
is difficult to ascertain but it seems unlikely that a spe-
cies which does not seem to form pairs and whose
males spawn with several females in a few days, would
normally behave in this manner.
X. ornatipinnis is thus a very interesting Tangan-
yikan cichlid which may provide us with even more
surprises. Its refined attractiveness and peaceful
behavior should help to establish its popularity among
aquarists in the years to come.
A brooding female Xenotilapia ornatipinnis.

18
 Ctenochromis benthicola (Matthes, 1962)
Mark Smith

A female Ctenochromis benthicola. This specimen was imported as “Orange Compressiceps”. Photographer unknown.

One of the most unusual and rare cichlids presently on its way to a natural extinction in the years to fol-
known from Lake Tanganyika is Ctenochromis low. On the other hand some species naturally have
benthicola. The specimen in the accompanying pho- small numbers because that keeps them in balance with
tograph is a female from the northern end of the lake. their biotope.
This individual somehow made its way into the U.S.A. C. benthicola is also endowed with enlarged sen-
in the early 70’s under the name of “Orange sory pits in the head, a feature shared with members
Compressiceps”. Whether or not more of these cich- of the genus Trematocara, which also inhabit great
lids were ever brought out of the lake for the aquarium depths. These enlarged pits probably enable these cich-
hobby is uncertain. lids to find their food in the relative absence of light.
Before Poll’s revision of the Tanganyikan cichlids C. benthicola was successfully bred by the
(1986) this species was included in the genus Brichards in the mid 70’s. Curiously details of its re-
Haplochromis. Now it is assigned to Ctenochromis, production were absent from their report, but since
which seems to be an odd genus since representatives only a very few juveniles were produced it may be
of it are found well outside of Lake Tanganyika and that C. benthicola is a mouthbrooder like all other
seem to be of a different lineage. haplochromines. The juveniles were brown at a length
Males of this species are a dark brown with a dark of about 5 cm and thus looked like their father. Some
blue overlay, while females are entirely orange with questions must arise from this bit of information. Did
just a few black specks on the sides of the body. This the spawn contain only males or do females start out
peculiar color scheme is unique among Tanganyikan looking like the brown males and then change color
cichlids. The brilliant orange color of the female would as they mature. Another possibility is that the orange
seem to make it more conspicuous. females are a kind of color morph, like the O and OB
The intense competition in the shallow waters of color morphs among Malawian cichlids. (According
Lake Tanganyika explain why this fish inhabits only to Mireille Schreyen from Fishes of Burundi, brown
the deeper waters. Its scarcity in the lake may also be females occur as well making it likely that the orange
due to the greater success of other cichlids causing C. individuals are indeed a polymorph. Ed.).
benthicola’s numbers to dwindle, indicating that it is

19
 Neolamprologus petricola (Poll, 1949)
Ad Konings

A wild caught male Neolamprologus petricola.

The name “Lamprologus Petricola” has been around colorless or greyish but never yellow. Large specimens
in the hobby for a long time. It was given to the yel- of N. petricola show a cranial gibbosity which has
low morphs of Neolamprologus mustax and seems still never been observed in N. modestus. The latter spe-
to be in use. Interestingly, Neolamprologus petricola cies has thicker lips too, but the teeth on the pharyn-
resembles N. moorii and N. modestus more than N. geal jaws are similar and indicate a diet of hard inver-
mustax. The name petricola means “living among tebrates for both species. The two species may differ
rocks” and alludes to the assumption that this cichlid in habitat preference as N. petricola is also found at
is restricted to rocky habitats. N. petricola inhabits deeper levels whereas N. modestus seems to be re-
the intermediate biotopes along the southwestern coast stricted to the shallow regions of the sediment-rich
of the lake. It has never been observed in Zambia. Al- intermediate biotope.
most all specimens exported were collected south of The first observations in the aquarium indicate that
Moba in Zaïre, but it is found as far north as Cap Tembwe. N. petricola is a territorial cichlid which is difficult to
It is very difficult to formulate a distinction between house with conspecifics in a small tank (less than 200
N. petricola and N. modestus, especially when semi- litres). The sexes are easily recognized by examining
adult. Both species have an elongated shape, but, in the vents and thus a single pair can be selected and
general, N. modestus has a shallower body. Both spe- placed in a large tank. It is the female who starts dig-
cies have a dark brown coloration but under subopti- ging a nest under a rock and she tries to attract the
mal conditions both species show a much lighter pat- male to her nest.
tern. N. modems, in such circumstances, has a much N. petricola grows to a size of about 13 cm and it
yellower tinge than the beige-greyish colored N. seems to attain the deeper body and the gibbosity on
petricola. Submissive coloration of the latter species the head only at adult size. As in N. moorii the female
shows grey, broad vertical bars on an almost white is slightly larger than the male and she is able to pro-
background. I have never seen the forms of N. duce large spawns. A spawn from a relatively young
modestus from areas other than the deep south of the female numbered more than 300 fry and it seems that
lake and these types are characterized by bright yel- fully grown females are able to produce more than 500
low pectoral fins. The pectorals of N. petricola are eggs per spawn. The juveniles have a light grey color.

20
 Neolamprologus leloupi (Poll, 1956)
Ad Konings

Neolamprologus leloupi at Kapampa.

In the first volume of the “Cichlids Yearbook” I have

written an entry about a Neolamprologus with a yel-
low dorsal fin which I named N. leloupi. At that time
I had not observed this species in its natural environ-
ment and agreed with Brichard (”Cichlids and all the
other fish of Lake Tanganyika”, 1990: 537), who had
surveyed the entire Zaïrean coast, that N. caudo-
punctatus is a synonym of N. leloupi.
In September 1991, Martin Geerts and I made a
short visit to the southern Zaïrean part of the lake and
also dived near a village named Kapampa. In this area
the distributions of N. leloupi and of N. caudopunctatus Neolamprologus leloupi at Kanoni, Zaïre.
overlap. This is also the only location where N.
caudopunctatus has a bright yellow dorsal. This may
have been the sole reason why these two species do
not hybridize. N. leloupi is distributed north from
Kapampa up to Cape Tembwe whereas N. caudo-
punctatus is found south from this region (including
the entire Zambian shoreline of the lake).
N. leloupi occurs in several population which dif-
fer slightly in the markings in the fins. There is no
difference between its behaviour and that of N.
caudopunctatus, and at Kapampa groups of both spe-
cies are found side by side in the intermediate habitat.
They occur from the shallows to a depth of about 25 m.
Neolamprologus caudopunctatus at Kapampa.

21
 Neolamprologus sp. “Ubwari Buescheri”
Mark Smith

One of the first exported Neolamprologus sp. “Ubwari Buescheri”. Photo by Mark Smith.

The existence of this undescribed Neolampmlogus sp. present (Chituta Bay population). N. sp. “Ubwari
“Ubwari Buescheri” was first made known photo- Buescheri” does not show such geographical varia-
graphically to the aquarium world in Axelrod’s tion. In fact the upper horizontal stripe of this species
Aquarium Atlas (1985; first ed. p 511, plate # 349) is at a noticeably different angle than in N. buescheri.
where it was wrongly identified as Lamprologus Furthermore N. sp. “Ubwari Buescheri” shows cra-
buescheri. According to Pierre Brichard (1990), his nial gibbosity at a small size, 6-7 cm TL, suggesting
son Thierry discovered this lamprologine at the Ubwari that 6-7 cm constitutes the adult size for this fish. This
Peninsula in February of 1984. Konings (1988) re- cranial gibbosity seems to be lacking in N. buescheri
ports that a specimen of this species was found to- for the most part. Lastly, from superficial observations,
gether with a collection of other cichlids from Burundi, the trailing ends of the unpaired fins of N. sp. “Ubwari
which could indicate that it is also found in this part Buescheri” tend to be finer or threadlike in contrast to
of the lake. (It has not yet been observed in Burundi the thicker fin-tips seen in N. buescheri. All these ob-
waters. See next page for the discussion of a similar servations plus further scientific analysis of this
species from the Kavala Islands. Ed.) lamprologine will undoubtedly show that these two
There are a number of differences between this spe- cichlids are not conspecific.
cies and N. buescheri. The “Ubwari Buescheri” has N. sp. “Ubwari Buescheri” was found at a depth range
smaller, more numerous scales than N. buescheri from of 15 to 25 metres, a range that basically matches that of
the southern part of the lake. The horizontal striping N. buescheri. It has also proven itself a territorial
in N. sp. “Ubwari Buescheri” is much narrower – it aquarium resident; like N. buescheri it forms pairs and
can at times even fade completely in some aquarium defends a small section of the aquarium as a territory.
specimens – than that of N. buescheri, The basic pat- The five specimens, two females and three males, we
tern of N. buescheri consists of two horizontal stripes keep in our tanks have yet to reproduce, but it is prob-
and several vertical stripes. Depending on the loca- ably safe to say that they spawn in the same fashion as
tion either the vertical striping (Gombi population) or similar-sized, rock-dwelling lamprologines do. The finely
the horizontal striping (Kachese population) is pre- elongated fins make N. sp. “Ubwari Buescheri” a grace-
dominant or else a combination of both patterns is ful addition to the spectrum of Tanganyikan cichlids.

22
 Neolamprologus sp. “Kavalla”
Ad Konings

A male Neolamprologus sp. “Kavalla” photographed at Milima Island at a depth of about 15 m.

Neolamprologus sp. “Kavalla” closely resembles N. mens, probably females and sub-adults, rarely display
sp. “Ubwari Buescheri” (see previous page) in shape the grey-blue colour. These individuals show a basic
and in some of its pigmentation pattern but grows to a pattern of two horizontal lines in which they resemble
much larger mature size. The maximum size of N. sp. N. sp. “Ubwari Buescheri”.
“Kavalla” is estimated at 12 cm (no specimens were All adult specimens have a dark grey-brown patch
collected for closer examination). on the lower part of the head, which is most intensely
I have observed this attractive cichlid at Milima Is- coloured near the chin where it abruptly disappears,
land, one of the islands in the Kavalla archipelago north leaving a pure white chin. Although the observation
of Kalemie on the Zaïrean shores of the lake. Its that adult size and coloration differ from those of N.
biotope consists of large, rounded boulders which sp. “Ubwari Buescheri”, it is still possible that we are
showed at the time of my visit (September 1991) just dealing with two populations of one species. Not un-
a thin layer of biocover. The coast around the island til the rocky regions between Ubwari and Kavalla have
shelves gradually to a depth of about 30 m where the been investigated and no intermediate Lamprologines
habitat becomes predominantly sandy. N. sp. have been found may we conclude that these two popu-
“Kavalla” is seen from 10 m on to the edge of the lations belong to different species. The basic pigmen-
rocky part at about 30 m. It normally occurs solitary tation pattern, i.e. the two horizontal lines on the body,
and is not common. One would expect that, like N. is seen in a small number of cichlids belonging to
buescheri, males at least would have territories, but Neolamprologus as well as Lampmlogus. It seems to
territorial individuals were not seen. It seems that N. be a characteristic of cichlids which are rare and usu-
sp. “Kavalla” roams through the biotope in search of ally live at deep levels. Besides the two species men-
food which presumably consists of invertebrates. It tioned here there is also L. sp. “Zambia” (see photo in
was frequently seen swimming in and out of caves in “Tanganyika Cichlids” by Konings, 1988: 108) and
which it may find most of its food. In this respect it frightened specimens of N. sp. “Cygnus” (see photo
behaves more like N. leleupi than like N. buescheri. in “Cichlids Yearbook”, vel. 1, 1991: 11) having such
Only the larger individuals, presumably males, show a colour pattern.
a grey-blue colour on head and body. Smaller speci-

23
 Cyphotilapia frontosa (Boulenger, 1908)
Ad Konings

The race of Cyphotilapia frontosa at Kavalla, Zaïre (photo above) resembles that of Burundi.

It is characterized mainly by the dark blue colour on

the cheeks and the variable yellow coloration in the dor-
sal fin. The third variant exported from the lake comes
from Zambian waters. It is characterized by a brighter
blue coloration (in comparison with the northern races)
and an interorbital band. In the late eighties a very dense
population of C. frontosa was discovered near the Kavalla
archipelago in Zaïre. This variant resembles the Burundi
race but occurs in rather shallow water (up to 5 metres).
In 1990 Alain Gillot (Zaïre Cichlids) discovered a new
and exciting variant of the frontosa. This population is
Plecodus straeleni photographed in Burundi waters. characterized by the intense blue colour on all fins and
the mother of pearl on the hump and upper part of the
body in large specimens. Juveniles also show the attrac-
The first live specimens of Cyphotilapia frontosa were tive blue coloration seen in adults. There is, however,
exported by Fishes of Burundi in the late fifties. From one drawback; it occurs at deep to very deep levels. The
then on it has been a mainstay in the aquarium hobby shallowest point I have seen this blue frontosa was 25
and become widespread among aquarists. C. frontosa metres. Most specimens are collected at a depth between
has a lake-wide distribution and occurs in several distin- 40 and 60 metres and require three days to decompress.
guishable races. Initially only the populations of Burundi The blue variant is distributed on the rocky shores be-
were exploited. Then the Kigoma race was introduced tween M’Toto and Kapampa on the Zaïrean coast.
to the hobby. The latter variant shows one vertical band An interesting phenomenon is the occurrence of a
more on the body than the Burundi race and has thus scale-eating cichlid, Plecodus straeleni, which has an
become known as the Seven-Band Frontosa. In view of identical coloration to that of C. frontosa. It was re-
the other variants known to date it is better to refer to it ported (Brichard, 1978, 1989; Krüter, pers. comm.)
as C. frontosa (Kigoma). that P. straeleni uses its coloration as a camouflage

24
The blue race of Cyphotilapia frontosa is distributed along the southern part of the Zaïrean coast.

which enables it to mix with groups of C. frontosa

and stealthily scrape some scales from the flanks of
the surprised frontosas. I have never observed P.
straeleni attacking C. frontosa although I have seen
them among their schools. However, I have observed
P. straeleni attacking other cichlids, like Cyathophar-
ynx furcifer or even the much smaller Neolampmlogus
brichardi.
It has also been said that it may mimic not only C.
frontosa but also Neolamprologus sexfasciatus and
N. tretocephalus. There are several reasons why this
seems unlikely. First of all P. straeleni resembles C. Plecodus straeleni at Kapampa, Zaïre.
frontosa to such an extent that it is easily mistaken
for its model, even at close quarters. It goes to great
lengths to copy C. frontosa, so much so that the popu- combine this with the observation that P. straeleni at-
lation at Kapampa has a much bluer coloration (like tacks other cichlids than C. frontosa then it seems
C. frontosa) than that at Rutunga, Burundi (see pho- unlikely that P. straeleni needs its coloration to be
tographs). Secondly, in Burundi waters there are no able to exist predominantly on scales of C. frontosa.
N. sexfasciatus or N. tretocephalus, which would of C. frontosa roams about in the rocky biotope but it
course not exclude the theory that P. straeleni’s col- has never been seen hunting prey. Other species in-
our pattern could have developed in mimicry of those habiting the same biotope probably know its peaceful
species, but it would make the extreme resemblance manners and thus give P. straeleni, dressed in a
to C. frontosa at locations where these two species “sheepskin”, a good opportunity. Feigning to be a
are found inexplicable. Thirdly we could argue that good-natured C. frontosa, the scale-eater comfortably
other scale-eaters, especially the abundantly present closes in on its prey and before the victim recognizes
Perissodus microlepis, do not mimic their prey. If we the wolf it has lost some of its scales.

25
 Cyprichromis sp. “Leptosoma Jumbo”
Ad Konings

The yellow morph of Cyprichromis sp. “Leptosoma Jumbo” at Kitumba, Zaïre.

The genus Cyprichromis consists of several species The observation that a female may give birth to
which belong to the most colourful cichlids of Lake yellow as well as blue tailed males in captivity does
Tanganyika. Representatives of the genus are found not prove the natural cause of things. Such a female,
at most rocky regions around the lake but none of the e.g. originally a yellow-tailed “species”, may have
species has a lake-wide distribution. Only two spe- spawned with a blue-tailed male in the confines of an
cies are scientifically described but at least two others aquarium. If the colour of the tail constitutes the main
are known to occur. At some locations in Zambia (e.g. criterion for mate-recognition, such a female would
Mpulungu; René Krüter, pers. comm.) and Tanzania never have done so in the wild. Until unambiguous
(e.g. Malasa Island; Walter Dieckhoff, pers. comm.) experiments have been performed revealing the in-
two species of Cyprichromis are found sympatrically, heritance of the colour of the tail, the possibility of
but at most other locations it seems that only one spe- two sympatric species cannot be excluded.
cies of this genus occurs. However, they are usually Sometimes, when such “hybridization” experiments
found sympatrically with Paracyprichromis brieni and seem to have occurred in the wild, we may deduce
P. nigripinnis. false clues regarding the specific status of the differ-
It is generally known that in a single population of ent sympatric species. This may be the case with the
Cyprichromis males with a noticeably different col- population of Cyprichromis at Kitumba, Zaïre. In this
our pattern, especially in the tail, occur in the same population not only the colour of the tail differs but
school. I have suggested before (1988) that the blue- also the colour on the body. In fact there are three dif-
tailed individuals could in theory be a genetically dif- ferent morphs (or species?): there are completely yel-
ferent species from the yellow-tailed specimens. Their low coloured males and blue coloured males with yel-
behavioural preference for feeding from plankton low and blue tails.
above rocky substrates may have brought these two I could observe this school for only one hour but
theoretical species together as it brought Para- during this short period I gained the impression that
cyprichromis, which has a similar feeding behaviour the complete yellow individuals and the blue individu-
but is from a different ancestry, together with als represent two different species. As is usual among
Cyprichromis. Cyprichromis, males defend their three-dimensional

26
 territory in the open water and relate the boundaries
to the distance from neighbouring males. Yellow and
blue males were found mixed and among them large
numbers of females, which all looked identical, tried
to find a partner to spawn. Both types of males some-
times courted the same female but their territorial ag-
gression was always directed towards a male of the
same colour; i.e. yellow males would chase only other
yellow males while blue males (with blue or yellow
tails) chased only other blue males. Sometimes a chase
followed a route through the territory of a differently
coloured male!
Notice a small aberration in this male’s tailfin. After a while I noticed a few rare individuals which
looked like a cross between the two morphs. Although
I do not know what a cross between these two morphs
would look like, the colour patterns these individuals
showed did not indicate that a cross would yield simi-
lar looking offspring. The colour aberration I noticed
and photographed (see accompanying pictures) were
mainly a type of local mutation rather than a recurrent
pattern of a segregation of the parental colors. So there
was a mainly yellow individual with a blue blotch on
its body or vice versa, a blue male with a yellow spot.
Although it cannot be denied that the yellow males
are distinctly different from the blue ones, it is still
not clear whether females appreciate these differences
A male of the yellow morph with a broad blue segment.

The dorsal fin of the male of the blue morph (here with yellow tail) becomes almost white when he courts.

27
as well. The fact that I could find three aberrant indi- probably an insignificant part of the total population.
viduals among a few hundred males may indicate that If such contrastingly coloured males can belong to
they are not products of an extremely rare event. one species then how can we explain the existence of
These observations can be explained in several dif- two species of Cyprichromis at Malasa Island and
ferent ways of which true hybridization of two Mpulungu? The colour difference between the males
sympatric but distinct species is just one. The yellow of these two species is much less dramatic than in the
and blue males could represent different morphs of Kitumba population. Maybe the latter population can
one species and a female would spawn with all three teach us a lesson on speciation. It may turn out that
of them. The territorial colour of the males could be coloration is not the most important criterion in spe-
regulated by one or very few genes which could act cies recognition but that species specific odours com-
like a switch in a juvenile phase of the male: normally bined with behavioural and morphological characters
it would switch on either blue or yellow but in some play a major role instead.
individuals and at certain regions of the body the switch A possible advantage for Cyprichromis having dif-
could have been reversed. This could occur as a ferently coloured males which do not visually recog-
phenotypic event, independently from genetic factors. nize each other as belonging to the same species, is
If the males of this population are to be regarded as that their territories can be closer together without elic-
morphs of one species then they certainly present a iting constant territorial fights of similar looking,
remarkable case of polymorphism (see photographs neighbouring males. Indeed I frequently observed
for how different the two morphs are). If the two yellow and blue territorial males almost perfectly al-
morphs represent two species then the definition of a ternating in the water column, For the time being I
natural species (specific recognition under natural cir- regard the Kitumba population as belonging to one,
cumstances) seems to be in need of a revision because polymorphic species, namely Cyprichromis sp.
females apparently can mistake a male of the other “Leptosoma Jumbo”.
species as their true mate. Mary Bailey (pers. comm.),
however, suggests that misfertilisation could easily
occur in a mêlée, and that the odd males I saw were

The completely blue morph of Cyprichromis sp. “Leptosoma Jumba” from Kitumba.

28
 Paracyprichromis brieni (Poll, 1948)
Ad Konings

A wildcaught male Paracyprichromis brieni collected at Kitumba, Zaïre. Note the spots in the trailing part of the dorsal fin.

It is a geographical variant of P. brieni which is

distributed along the Zaïrean coast between Kitumba
and Moba. It has a rather wide depth-range since indi-
viduals were observed between 5 and 35 metres.
Since its introduction it has been bred in captivity,
but reports concerning a spawning are not yet avail-
able. Courting behavior differs slightly in some points
from that of P. nigripinnis. Males occupy territories,
like P, nigripinnis males, but these are not necessarily
alongside the vertical face of a rock or at the side of
the tank. Females stay in the open water as well and
form schools. Male P. nigripinnis court females by
A female Paracyprichromis brieni (Velifer).
displaying themselves with all fins erect, followed by
leading the female to the centre of its territory while
When the first specimens of this pretty little cichlid – swimming in an undulating fashion.
with the trade name “Cyprichromis Velifer” – arrived The courting behavior of P. brieni (Velifer) differs
in western aquarists’ tanks it was immediately thought in that some males do not lead the females while
that the individuals with the attractive black and white vigourously quivering their bodies. The male starts
dorsal fins were males and the ones with the plain fins courting with a display of all its fins accompanied by
were females. Of course, after acclimatization and with a slight quivering and sometimes sideways jerking of
observations made under less stressful conditions it the body. Meanwhile he inches his way towards the
became apparent that it is the female which is adorned female and, at close range, gives her a sudden blow
with the black and white dorsal fin. Due to the lack of with his tail. Males P. brieni (Velifer) are character-
yellow coloration on the tips of the ventral fins this ized by a black spot in the middle of the tailfin. As far
species is placed in the genus Paracyprichromis rather as is known this is the only geographical variant of P.
than Cyprichromis. brieni with such a feature.

29
 MALAWIAN
CICHLIDS

The genus Cynotilapia Regan 1922

Dr. Andreas Spreinat

The main typical feature of the species of the genus plankton rather than of Aufwuchs. The unicuspid,
Cynotilapia is unicuspid teeth, spaced relatively far widely spaced, teeth are better suited to the capture of
apart and resembling the teeth of a predator. Most other small planktonic invertebrates (zooplankton) than to
morphological characteristics resemble those of the combing the algae of the biocover.
members of the Pseudotropheus zebra complex to such The type species, C. afra, was described in 1893
an extent that without knowledge of the tooth struc- by the ichthyologist Albert Günther, who worked at
ture, they can easily be confussd with these mbuna. In the British Museum (Natural History). In 1976, the
nature another feature can be observed that further second species in this genus, C. axelrodi, was described
differentiates Cynotilapia from Pseudotropheus: in by Burgess. Tony Ribbink and his co-workers, a South
contrast to most other mbuna, Cynotilapia, except for African team of ichthyologists, concluded, after the
territorial males, forages in the water column several investigation of many rocky habitats along the shores
feet above the rocky substrate. Their food consists of of Lake Malawi, that a further 8 species of Cynotilapia

A male Cynotilapia afra in the rocky habitat around Chitandi Island; note its teeth!. Photos by A. Spreinat.

30
exist (Ribbink et al., 1983). These species were not number of males with black dorsal fins decreased to-
formally described but given provisional names. wards the south, whereas males with white or yellow
Probably even more species exist. Some time ago dorsal fins became more abundant there.
specimens from a population of Cynotilapia were Among aquarists it was previously common prac-
imported under the trade name “Jaro Afra”. These cich- tice to differentiate between Cynotilapia and
lids, collected at Jalo Reef near Nkhota Kota, belong Pseudotropheus by looking at the vertical bars on the
to a scientifically undescribed species provisionally body. These continued into the dorsal fin in Cynotilapia
called C. sp. “Jalo”. and not in Pseudotropheus. Owing to the increased
Among aquarists C. afra is the best known species. knowledge of species and geographical variants this
I am not aware when it was exported for the first time, criterion has been invalidated, as some species of
but it must have been in the late sixties or early seven- Cynotilapia are known to have entirely white dorsal
ties. C. afra is found in the northern half of the lake. fins while some members of the Pseudotropheus ze-
Likoma, Chizumulu and the rocky shores between bra complex have vertical barring extending into this
Chirombo Point and Ngara on the West coast are fin. Given the considerable variation known to occur
known to harbour this mbuna. within a population of C. afra, it is difficult to assess
The variation in the colour pattern of C. afra is note- whether in other populations we are dealing with a
worthy. On the one hand populations show geographi- geographical variant or with a different species. Com-
cal variation in male breeding coloration, and on the plicated and elaborate investigations are necessary in
other hand variations exist within a single population. order to make a final pronouncement.
These variations are not just subtle changes in the col- By comparison the recognition of C. sp. “Mbamba”
our pattern but include conspicuous differences in the as a valid species is relatively simple. C. sp. “Mbamba”
nuptial coloration of males, especially the coloration lives sympatrically with C. afra at most locations al-
of the dorsal fin. Ribbink and his coworkers investi- beit at a generally deeper level. Also C. sp. “Mbamba”
gated the differences in the coloration of the dorsal shows a considerable variation in male breeding col-
fins of males in the populations inhabiting the north- oration, geographically as well as within a single popu-
ern shores of Likoma Island. They found that the lation. Some individuals are almost completely black,

A male with a black dorsal. Is this a geographical variant of Cynotilapia afra or a different species?

31
i.e. the black bars are so broad that they cover most of found. Ribbink and his coworkers (1983) and several
the flanks. Other specimens, however, have a colour diving aquarists found C. axelrodi to be restricted to
pattern not unlike that of Pseudotropheus zebra. Some the western coast between Chirombo Point and Li-
males have a yellow, and others a light blue, blaze on on’s Cove. Another species, which belongs to
top of the head. This part of the head is also the sim- Pseudotropheus because of its bicuspid teeth, collected
plest feature by which one can discriminate between at Likoma Island and also named “Kingsizei”, further
C. afra and C. sp. “Mbamba”. C. afra has a black in- confused the identity of C. axelrodi.
terorbital bar (from eye to eye) and a bar across the Not only does C. axelrodi differ from the other spe-
upper part of the head. This second bar, which runs cies of the genus in its almost nondescript, light blue
between the upper parts of the gillcovers, is lacking in coloration, but it also has an elongated shape. The
C. sp. “Mbamba”. Sometimes a faint trace of the bar population which I was able to observe north of Nkhata
is visible but the background colour, yellow or light Bay inhabited shallow water over a mixed bottom at a
blue, is clearly visible giving the fish the appearance depth of 4-6 metres. Males behaved territorially and
of having a coloured cap. defended areas with a diameter of about one meter.
Unfortunately C. sp. “Mbamba” was exported only The only other Cynotilapia which is elongated like C.
sporadically although it would make a very attractive axelrodi, is the scientifically undescribed C. sp. “Lion”.
addition to the mainstays of the hobby. C. axelrodi, This cichlid, with its trade name “Lions Cove Afra”,
however, is a frequently exported member of the ge- is, as its name indicates, collected near Lion’s Cove
nus. It is usually exported under the trade name on the northwestern shore. C. sp. “Lion” also prefers
“Kingsizei”. The first specimens were exported by a mixed sandy-rocky bottom in shallow water. It is
Peter Davies in 1973. For a long time the identity of rarely found in waters deeper than 20 metres. Males
C. axelrodi was uncertain, probably because Burgess, excavate small caves into which females are led in
who described the species in 1976, gave as type local- order to spawn. Like many other mbuna, C. sp. “Lion”
ity the southern part of the lake (Burgess, 1976: 41). tends to lose its elongated body-shape when fed copi-
This was the location from which the exporter oper- ously. Such specimens are often as deep-bodied as wild
ated but not where this ’species was subsequently C. afra.

A male Cynotilapia sp. “Mbamba” at Chitande Island.

32
 The distribution of C. sp. “Chinyankwazi” is lim- of Mbuna from Lake Malawi (Pisces: Cichlidae).
ited to the two small islets, Chinyankwazi and Trop. Fish Hobbyist 24, Oct. pp 37-44.
Chinyamwezi, in the southeastern arm of the lake. GÜNTHER, A. (1893) Second report on the reptiles,
(Recently a population has been discovered at batrachians and fishes transmitted by Mr. H. H.
Chinyamwezi Reef, Ed.). These two islets belong to Johnson, C. B., from British Central Africa. Proc.
the National Park and are forbidden grounds for the zool, Soc. Lond. pp 616-628.
collection of ornamental fish. The coloration of C. sp. RIBBINK, AJ., B.A. MARSH, A.C. MARSH, A.C. RIBBINK,
“Chinyankwazi” is almost identical to that of and B.J. SHARP (1983) A preliminary survey of the
Pseudotropheus zebra. Interestingly, P. zebra is not cichlid fishes of rocky habitats in Lake Malawi. S.
present at these two islets. It would be worth the ef- Afr. J. Zool 18 (3), pp 149-310.
fort to investigate how the niche of C. sp. “Chin-
yankwazi” compares with that of P. zebra. It is possi-
ble that C. sp. ’Chinyankwazi” has been exported in
earlier times, when collection at these two islands was
still allowed. Maybe some specimens of this mbuna
are still around in the hobby, possibly under the name
P. zebra or as an unidentified species.
The other species, for which there is no space for
photographs, will be discussed for completeness.
Ribbink and his coworkers identified a rare species at
Ndumbi Rocks, Likoma Island with the name C. sp.
“Ndumbi”. It lives in secluded caves in shallow wa-
ter. Its coloration is almost entirely black.
C. sp. “Maleri” is known only from the Maleri Is-
lands (Ribbink et al., 1983). It has a dark blue body
with black vertical bars. The Mbenji Islands harbour C. axelrodi 2 km north of Nkhata Bay.
two species of this genus, C. sp. “Yellow Dorsal” and
C. sp. “Black Dorsal”. The main characteristics of the
male breeding coloration are used in their unofficial
names. C. sp. “Jalo” was mentioned earlier in this ar-
ticle.
The maintenance of Cynotilapia is not much dif-
ferent from that of other mbuna. Besides a slightly
alkaline pH-value of the water and a modest feeding
regime, the size of the tank and its decoration is an
important factor. Under the right conditions and with
a restricted but varied diet Cynotilapia behaves as a
lively and robust mbuna. As with many other mbuna,
territorial Cynotilapia can be aggressive towards A male Cynotilapia sp. “Lion” at Lions Cove.
conspecifics as well as other species.
Most of the imported species can be bred in captiv-
ity. However, little has been reported about the inher-
itance of the different colour patterns in C. afra and
C. sp. “Mbamba”, the two species with the variable
coloration. There is thus an opportunity, especially for
aquarists, to discover some interesting facts about these
fascinating mbuna.

References

B URGESS , W. E.. (1976) Studies on the family

Cichlidae: 7. Cynotilapia axelrodi, a new species Cynotilapia sp. “Chinyankwazi” at Chinyankwazi Island.

33
 The Protomelas taeniolatus-complex
Ad Konings

The race of Protomelas taeniolatus at Chinyankwazi Island is better known as “Haplochromis Fire Blue”.

Protomelas taenioIatus is one of the most frequently are dealing with two different species which can be found
exported non-mbuna from Lake Malawi. In the aquaristic sympatrically. Ten years ago the scientific identification
trade it is better known as Haplochromis Steveni. Dur- of these two species was still unclear. It was Tony Ribbink
ing the lake’s evolution many geographical races have and his coworkers (1983) who gave us important new
developed. Several localities are regularly fished for these information about these two species. They pointed to
variants which have all been given separate names. A the fact that these two species behave differently.
few aquarium favourites received names like “Steveni Protomelas taeniolatus, the most common of the two,
Tiger”, ’Red Empress”, or “Fire Blue”, while others were feeds on the biocover in the upper regions of the rocky
named for the locality where they were collected, e.g. habitat. Its food consists mainly of algae. Protomelas
“Steveni Mbenji” or “Steveni Maleri”. fenestratus, the “Steveni Thick Bars”, lives in the deeper
It was known to the fish-collectors that one location regions of the rocky biotope and blows away the sedi-
could harbour two varieties of “Steveni”. “Steveni Ti- ment on the rocks to reveal its prey, which consists mostly
ger” and “Steveni Thick Bars” were both caught at of insect larvae and crustaceans.
Likoma and Chizumulu Island. Although they have dif- Even though in shape and appearance these two spe-
ferent preferences concerning their habitat, both Stevenis cies are sometimes difficult to distinguish, by observing
could be seen together at some locations. “Steveni Ti- their feeding behaviour one can easily identify them.
ger”, now identified as Protomelas taeniolatus, has thin Ribbink et al. (1983) further noted that the colour pat-
vertical bars on a silvery body, whereas “Steveni Thick tern of P. fenestratus consists of conspicuous vertical
Bars” has broad vertical bars. The nuptial coloration of bars whereas that of P. taeniolatus has more horizontal
the male differs too. A “Steveni Tiger” male has an in- elements. Now, with knowledge of behaviour and col-
tense blue on the head and upper part of the body and an oration it must be easy to identify all populations of these
intense orange-yellow on the flanks. A “Steveni Thick cichlids in the lake. Initially, it seemed to work. I could
Bars” male has a light blue breeding coloration and a (I thought) identify every population even though I found
yellow throat. Its colours are much less intense com- that the pattern of markings, especially in P. taeniolatus,
pared to the “Steveni Tiger”. varied considerably (which was also mentioned by
From the previous description it must be clear that we Ribbink et al.).

34
 After more than 250 hours of diving experience in the lect it. Since the latter specimen showed distinct vertical
lake, the identification of these species looks less sim- bars I provisionally identified it as P. fenestratus although
ple. P. taeniolatus is very common and territorial males I didn’t see it blowing in the sand. I examined one speci-
are found on almost every rocky shore. P. fenestratus is men of the black-bellied Steveni and could not find any
much rarer but can be found at most rocky areas as well. distinction between it and P. taeniolatus. I Therefore
Breeding males are not common. They are weakly terri- thought it to be a geographical race of the latter species.
torial and usually construct a shallow nest in the sand In November 1990 I revisited the northwestern coast
alongside a rock. At Hora Mhango, on the northwestern and found black-bellied Stevenis at most locations, al-
coast of the lake, P. fenestratus was found to defend a beit in very low numbers. Furthermore I found a (blow-
territory on top of a rock, like P. taeniolatus does. I also ing) P. fenestratus at Kande Island which had a heavily
found two other species which blow in the substrate as barred colour pattern, unlike the one I had seen before.
part of their feeding behavior. One of these, P. My conclusion is that there are at least three species in
pleurotaenia, lives in the shallow sandy and vegetated the P. taeniolatus complex. All of these species can be
areas and does not show geographical variation. Its pat- found sympatrically. The black-bellied Steveni is com-
tern consists of two thin, horizontal lines on a silvery mon only at Kande and Mphandikucha Islands. Its col-
body. The other species is closely related to Placido- oration seems to be quite constant along the northwest-
chromis johnstoni and therefore easily distinguished from ern shores. Territorial males defend a nest on a rock or
P. fenestratus. As far as I know, a cichlid with a small on the sand beside a rock.
mouth, vertical barring, and blowing in the sediment is a The rocky areas around the Nsinje river delta on the
sure identification for P. fenestratus. east coast of Lake Malawi provided me with another
In May 1989 I found at Kande Island (and at problem. In this area the so-called “Steveni Eastern” is
Mphandikucha Island) a Steveni-type cichlid which had collected. At first, I had identified this cichlid as P.
the lower half of the body completely black. I photo- taeniolatus (Konings, 1989). During a later visit I found,
graphed another Steveni-type as well but could not col- north of the river, another Steveni-type. The males were

The algal-garden denotes the centre of the territory of this The “Steveni Eastern” is a race of Protomels taeniolatus.
male P. taeniolatus (Thumbi West Island).

A male Protomelas sp. “Steveni Blue Black” (Masinje). A Protomelas sp. “Steveni Black-Belly” at Mdoka.

35
completely blue (see photo) and stayed in the upper re- could confirm this idea, but the shape of the short snout
gions of the habitat. At the same time, at deeper levels, I resembles that of the northern species. For the time be-
observed P. fenestratus females blowing away in the sand. ing, however, we will refer to the blue Steveni as
I then concluded that the “Steveni Eastern” must be P. Protomelas sp. “Steveni Blue Black”.
fenestratus, as I found males at deeper levels than the A reef north of Chizumulu Island, named Taiwan Reef,
all-blue Steveni. Moreover I saw a male “Steveni East- harbours only one member of the P. taeniolatus com-
ern” displaying for such a sand-blowing female. My only plex. The reef consists of large rocks which meet the
reservation at that time was that the females, which were sandy bottom at great depths. The depth and strong cur-
normally shipped with “Steveni Eastern”, didn’t show rent, which is commonly observed around the reef, make
much vertical barring whereas the sediment-blowing the rocks virtually devoid of sediment. The Steveni at
females had distinct vertical bars. I never questioned the Taiwan has very broad, vertical bars but I have not seen
fishermen about this matter, which I should have done. them blowing into the biocover. If it were P. fenestratus
In a later publication (1990) I showed the “Steveni East-
ern” as P. fenestratus.
During my third visit to the area I found territorial
males of P. fenestratus which looked like anything but
“Steveni Eastern”. The coloration of these males is very
much like that of “Steveni Thick Bars” at Likoma and
Chizumulu Islands. So there are three species at the east-
ern shores as well. “Steveni Eastern” is a race of P.
taeniolatus in which the females have few markings on
a silvery body. The P. fenestratus has never been ex-
ported and thus lacks a trade name.
The third species, in which the males are blue, could
in fact be closely related to the black-bellied Steveni from
the northwestern shores. I have not seen females which A female with an unusual colour pattern but identified as
P. fenestratus because of her particular feeding technique.

A male Protomelas fenestratus at its shallow nest in the sand (Nsinje River Outlet).

36
this could be explained by there being no sediment to in the upper 10 metres at any rocky coast I have vis-
blow in (P. fenestratus hardly ever blows in the gravel in ited. Most of the year there is a heavy current around
an aquarium). After examining several specimens I must Taiwan Reef, sometimes so strong that it is difficult
conclude that the “Steveni Taiwan” is not a race of P. to swim against, even with fins! The only species that
fenestratus. Its anatomy is closer to that of P. taeniolatus braves such a current is Pseudotropheus saulosi. P.
but I now think it is better to regard it as a different spe- sp. “Steveni Taiwan” may have been forced by the
cies altogether. I don’t like to split up species-complexes current to dwell in deeper regions. P. taeniolatus and
into many different species, because, although it seems the “Steveni Taiwan” may have the same ancestor but
to solve a taxonomic problem, it is like turning your back due to the complete isolation from all other popula-
on the really interesting questions like variation, tions it has developed into a different species, adapted
speciation, and evolution. to its specific environment. Its relatively large eye
Taiwan Reef may be the only isolated place in the could therefore be an adaptation to the depth at which
lake that has harboured a viable population of cich- it lives.
lids, even during the low water stands. We may thus
expect to find old species at this reef. Species that References
were probably never in direct contact with their sib-
ling species elsewhere in the lake. Speciation may KONINGS, A. (1989) Malawi cichlids in their natural
therefore have come to a stasis long ago. There are habitat. Verduijn Cichlids, Zevenhuizen, Nether-
not many purely rock-dwelling cichlids at the reef; lands
there is only one species of the Pseudotropheus ze- KONINGS, A. (1990) Book of cichlids and all the other
bra-complex, one from the Ps. tropheops-complex, fishes of Lake Malawi. TFH Publications, Neptune,
two from the Ps. elongatus-complex; there is only one USA.
Melanochromis and there are no species from the ge- RIBBINK, AJ., B.A. MARSH, A.C. MARSH, A.C. RIBBINK,
nus Labidochromis. Protomelas sp. “Steveni Taiwan” and B.J. SHARP (1983) A preliminary survey of the
behaves as P. taeniolatus but males defend their nests cichlid fishes of rocky habitats in Lake Malawi. S.
at depths below 20 metres. P. taeniolatus lives mostly Afr. J. Zool 18 (3), pp 149-310.

Protomelas sp. “Steveni Taiwan” is easily the most beautiful species of the P. taeniolatus-complex.

37
 Tramitichromis lituris (Trewavas, 1931)
Ad Konings

A territorial male Tramitichromis lituris (Mdoka).

The genus Tramitichromis is characterised by the pe- the rakers and be carried to the outside while the lighter
culiar shape of the lower pharyngeal bone. The teeth material, including invertebrates and algae, remains
on this bone are all slender and long. The teeth situ- inside the mouth. The strength of the rakers is prob-
ated at the front are the longest, which is an unusual ably needed to withstand the abrasive action of the
feature. These long anterior teeth are further charac- sand. If the rakers were higher, the separating effect
terised by their long tips which are bent backwards. might be more effective. The need to keep the tips of
In most other species the anterior teeth are small and the pharyngeal teeth in the same plane as those of the
their tips point forward. The reason for this particular gill-rakers could have led to the situation in
development in Tramitichromis is not clear, but most Tramitichromis where the anterior teeth are relatively
likely its origin lies in the feeding behaviour or in the longer than in other cichlids. The anterior pharyngeal
type of food. Another feature of Tramitichromis is the teeth of e.g. Lethrinops leptodon are not long and are
downward projecting blade of the lower pharyngeal below the level of the raker-tips.
bone. The upper edge of this blade runs horizontally I recognize five, possibly six species, in Tramiti-
in most cichlids but in Tramitichromis it projects down- chromis. Four species are described: T. brevis, T.
wards (more than 50’ in T. variabilis). variabilis, T. lituris, and T. trilineatus. Because of its
The lower gill-rakers in Tramitichromis are short different colour pattern I regard intermedius – placed
and robust, as in other sand-sifting Lethrinops, e.g. L. in Tramitichromis by Eccles & Trewavas (1989) – as
leptodon, L. macrochir. The first two rakers are usu- a member of the genus Trematocranus.
ally no more than small knobs on the gill-arch. The I have never been able to examine specimens of T.
central three to five rakers on the lower arches are trilineatus, so all information given here relates to the
wide and much larger. The tips of the broad rakers first three species. T. brevis was exported as
together form an almost horizontal platform at the ’Lethrinops Variabilis” or as ’Lethrinops Chizumulu”
bottom of the buccal cavity. The kind of grid they form and is easily recognized by its small adult size (about
may separate the heavier material from the lighter 14 cm) and by the prominent diagonal stripe on its
when a mouthful of sand and sediment is taken into flank.
the mouth. The heavier sand may thus sink between The type material of T. variabilis consists, in my

38
A female Tramitichromis lituris (Mdoka). A male T. brevis at its cave-crater nest. (Chizumulu).

A cave-crater nest of Tramitichromis lituris. A Tramitichromis lituris nest on top of a rock.

“Lethrinops Red Flush”, possibly Tramitichromis variabilis, is collected in Senga Bay.

39
The lower pharyngeal jaw of Tramitichromis lituris. A T. variabilis-type lower pharyngeal bone; note the
steeply inclined anterior blade. Photos by Gertrud Dudin.

opinion, of at least two different species. Specimens Not only N. eucinostomus but also T. lituris males had
of one of these are exported as “Lethrinops Red Flush” made, their nests on top of rocks! Such unusual be-
and they were all collected in Senga Bay. This species haviour in utaka can be explained by the observation
has only vague markings on the body. Trewavas (1931) that its females feed on plankton in the open water
reports that the northern races (species?) have (mostly) and the males are thus closer to the females (even if
a diagonal stripe. Such specimens have been collected they have to carry the sand up the rock). Lethrinops
around Likoma and Chizumulu Island. forages in the sand and males building a nest on top of
T. lituris has recently been observed in its natural rocks would diminish their chances of mating. At
biotope but has never, to my knowledge, been ex- Mdoka, however, I found females several feet above
ported. A species which I previously identified as T. the substrate and at first I presumed that these were
lituris, is probably an undescribed, but closely related attracted by the males that had their territories on the
species (Konings, 1990). They came from Senga Bay rocks. When I examined a few preserved specimens I
but have, since their first importation, never been seen found their long guts (2.5 times standard length) com-
again. pletely filled with phytoplankton. Hence it is possible
I have reported on the breeding behaviour of T. that the territorial males have adapted themselves to
brevis (Konings, 1990) and can add only that breed- the situation where females feed in the open water
ing was also observed in November. Since Stuart Grant column due to a plankton-bloom.
collected males in breeding colour in April, it may be It is still remarkable that a bottom-feeding species,
possible that T. brevis breeds throughout the year. The which apparently needs the protection (during spawn-
main feature of the male’s nest is a small stone under ing) of a cave-crater nest, switches to an open nest-
(and around) which a semicircular rim of sand (ox- type a few metres above the floor. I have observed
bow) is deposited (see photo), the so-called cave-cra- females accepting the alternative nest and spawning
ter nest. The female lays her eggs as far as possible with the occupant. It is not likely that T. lituris breeds
under the small stone. throughout the year as this species was not seen (at
A breeding colony of T. lituris was observed near the same location) in May, 1989.
Mdoka at the end of November. Most members of the
school were found between a depth of 7 to 15 metres. References
Territorial males had constructed a cave-crater nest.
Similar nests are constructed by T. brevis although they ECCLES, DH. & E. TREWAVAS (1989) Malawian cich-
use smaller stones. The breeding colony of T. lituris lids. The classification of some Haplochromine gen-
was mixed with a breeding colony of Nyassachromis era. Lake Fish Movies, Herten, Germany.
eucinostomus the males of which build shallow nests KONINGS, A. (1990) Book of cichlids and all the other
in the sand when at deeper (than 7 m) levels. It is fishes of Lake Malawi. TFH Publications, Neptune,
known with regard to N. eucinostomus that some males New Jersey.
construct a sand-nest on top of rocks whenever there TREWAVAS, E. (1931) A revision of the cichlid fishes of
is a lack of sites on the bottom. The colony at Mdoka the genus Lethrinops, Regan. Ann. Mag. nat. Hist.
had some males defending their nest on top of a rock. (10), 7; pp. 133-153.

40
 Stigmatochromis sp. “Modestus Makokola”
Ad Konings

A wildcaught male Stigmatochromis sp. “Modestus Makokola”.

Stigmatochromis sp. “Modestus Makokola” has been modestus.

exported recently, albeit in low numbers. As its name There is also a difference in behaviour. S. modestus
implies, it is collected at the Makokola Reef, south of lives in the rocky habitat where it usually hides in the
Boadzulu Island in the southeastern arm of the lake. It dark recesses. The dark brown coloration blends well
has also been observed at Boadzulu Island. with the environment and provides a splendid camou-
S. sp. “Modestus Makokola” differs from S. flage. It is a typical ambush hunter, its prey being
modestus in several anatomical features. Its average mainly small mbuna. Only during the breeding period
adult size ranges between 16 and 20 cm. Although S. do males stake out a territory in the rocky biotope,
modestus can grow to a size of about 25 cm, most often under an overhanging rocky ledge. Territorial
adult specimens have a length between 14 and 18 cm. males gather in small colonies numbering between 5
The “Modestus Makokola” has a deeper body and the and 15 individuals.
three spots are more distinct than in S. modestus. The The few encounters I had with S. sp. “Modestus
usually dark brown body coloration of S. modestus Makokola” revealed a different behaviour in this
obscures its markings but when stressed, three small, predator. During all encounters (only males were ob-
round spots are visible. The first spot (suprapectoral) served) the fish was out in the open, not hiding among
in S. sp. “Modestus Makokola” is elongated and about the rocks. At Makokola Reef it seemed to be more
twice as long as in S. modestus. The body coloration of common near the intermediate habitat at a depth of
the “Modestus Makokola” is silvery rather than brown. about 35 meters. A few times it was seen cruising over
The teeth of S. sp. “Modestus Makokola” are uni- the sand, probably hunting for small, sand-dwelling
cuspid and not densely packed, which places it in cichlids which are abundant in that area. At Boadzulu
Stigmatochromis rather than Otopharynx. I did not Island it was seen once. There it was seen hunting
observe S. modestus at Makokola Reef or at Boadzulu over large rocks, behaving a little like a pursuit hunter.
Island, but they were present at Chinyamwezi and Because of its carnivorous appetite it is desirable to
Chinyankwazi Islands. Although they were not found accompany S. sp. “Modestus Makokola” with larger
together, I believe that the “Modestus Makokola” is a cichlids. Apart from that it is fairly easy to maintain in
different species and not a geographical variant of S. a community aquarium.

41
 Dimidiochromis strigatus (Regan, 1922)
Peter Baasch

A territorial male Dimidiochromis strigatus photographed in the author’s aquarium.

Dimidiochromis strigatus has a lake-wide distribution it a stouter appearance.

but it is infrequently collected. Observations in their Wild caught male D. strigatus start colouring up at
natural habitat are rare and are restricted to subadult a size between 15 and 20 cm. They develop into beau-
individuals with a length of about 12 cm. Reports about tifully coloured cichlids with a characteristic red spot
its feeding behaviour range from pursuit piscivore, or behind the gill cover and a red anal fin. Males defend
feeding on invertebrates, to vegetarian feeding on territories when they have assumed the breeding col-
aquatic plants. If one considers the habitat in which oration, and behave rather aggressively towards other
D. strigatus is usually found, i.e. the shallow sandy inhabitants of the aquarium. Therefore the tank should
areas, most of the alleged diets could indeed make up have a size of at least 150 cm, but 200 cm would be
D. strigatus’ dinner. The oblique mouth indicates that better. In the territory the male constructs a shallow
insects that have been fallen into the water may also crater with a diameter of about 50 cm and a depth of
form a part of its diet. The size of its mouth and the about 5 to 10 cm.
observations in the aquarium, however, suggest that it Spawning takes place in this crater-nest. The number
behaves like a ambush predator preferring live food of eggs laid is remarkable. Once one of my females
above plants. released 230 fry after the incubation period! Raising
D. strigatus is irregularly exported as “Haplo- the fry poses some problems. Besides nitrate-free
chromis Sunset”. Most exported specimens have a water, the fry also need close attention when feeding.
length of about 12 to 15 cm. The largest specimen They don’t know when to stop eating. Many casual-
ever caught measured 24.4 cm total length. D. strigatus ties may arise when too much food is given at one
is laterally compressed, has a rather deep body, and time. At a size of about 2.5 cm they seem to cope bet-
its lower jaw is longer than the upper. The horizontal ter with the food.
mid-lateral stripe on the body is a characteristic of all D. strigatus and D. compressiceps should be kept
species of Dimidiochromis. separated as hybridization between these two species
Its morphological features resemble those of D. has frequently been reported. Juveniles of the two spe-
compressiceps although its body is deeper and less cies are difficult to tell apart.
compressed than that of D. compressiceps, which gives

42
 Copadichromis boadzulu (Iles, 1960)
Peter Baasch

A courting male Copadichromis boadzulu; the female is partly visible in the background.

When one looks through the aquaristic publications its, this active cichlid – maximum size is about 14 cm
for mention of Haplochromis or Cyrtocara boadzulu – needs a rather large aquarium. Initially it may be a
one gets the impression that this species is one of the little panicky and thus care must be taken to cover the
best known of the Malawian cichlids. The photographs tank properly.
accompanying articles, however, show a different spe- The basic pigmentation pattern shows some verti-
cies to the one on this page. The reason is simple. Those cal barring but this is seen mainly in breeding males.
who have read the latest revision of Malawi cichlids The main characteristic is a horizontal mid-lateral line
(Eccles & Trewavas, 1989) and the latest aquaristic which runs from the third vertical bar to the caudal
literature (Konings, 1989, 1990) know that the cichlid peduncle. A second horizontal (dorso-lateral) stripe,
which has been sold for years under names like which is characteristic of Protomelas taeniolatus, is
“Boadzulu”, “Hinderi”, “Red Empress” or (better!) as entirely absent. Fully coloured males have a white-
“Namalenje” belongs to a geographical race of blue blaze on the head which continues as a white
Protomelas taeniolatus. This has left us with the ques- marginal band in the dorsal fin.
tion of what does C. boadzulu look like. Of great interest was the breeding behaviour dis-
The “real” C. boadzulu is, as far as I know, a rare played by a male in my tank. He constructed a turret
cichlid, small populations of which inhabit the sandy of sand with a height of about 25 cm! When placed in
regions in the southeastern arm of the lake. Until now another aquarium the male again made a sand-cone.
only two localities are known, one at Crocodile Rock Spawning has not yet been observed. The number
in 5 metre deep water and one at Makanjila Point where of fry released per spawn, however, is rather small
the sand is at a depth of about 15 metres. Sightings in and never consisted of more than 19 babies. The basic
the natural habitat are very rare; breeding individuals pigmentation pattern and the construction of a spawn-
were not seen. ing-cone may indicate that C. boadzulu belongs to
C. boadzulu belongs to the utaka group and feeds Nyassachromis, since most other utaka in Copadi-
predominantly on plankton in the open water. In cap- chromis have a pattern of spots and are not known to
tivity it eats all kinds of aquarium foods. Although build spawning-cones on the sand.
intraspecific aggression remains within tolerable lim-

43
 Buccochromis rhoadesii (Boulenger, 1908)
Dr. Andreas Spreinat

A male Buccochromis rhoadesii courting a female (in the background). Photo by A. Spreinat.

Only two of the seven species which are currently as- sign of the sexual dimorphism known from adults. At
signed to the genus Buccochromis have experienced a this stage they all had a yellow coloration which was
wide distribution among aquarists. B. lepturus was most intense on the ventral part of the body. At an age
imported several years ago under the tradename of almost two years and at a length of about 18 cm the
“Green Lepturus”. A second species was first imported largest specimens began to show signs of the male
under the trade name “Haplochromis Lepturus” and adult coloration. The first sign was a blue sheen on
later as “Yellow Lepturus”. Its scientific name was the skin between the eyes and mouth which progressed,
uncertain for a long time until the revision of the with age, to the anterior part of the head and, later on,
Malawian haplochromines (non-mbuna) by Eccles and over the entire body. The yellow coloration decreased
Trewavas (1989) was published. Then it became clear concomitantly with the appearance of the blue. When
that “Yellow Lepturus” had been described under the I could differentiate between the sexes, I took a pair
name B. rhoadesii. and placed them in a 3-metre aquarium, while I left
B. rhoadesii probably has a lake-wide distribution the other pair in the tank they grew up in.
as it has been caught at several localities around the The complete metamorphosis of the males took sev-
lake. Most of the exported specimens were collected eral months. The male in the photograph measures 27
at Likoma Island. Here young solitary individuals or cm while the females, of the same age, measure be-
small groups are regularly seen over mixed sand-rock tween 20 and 23 cm.
habitats but also in rocky areas. They are seen in shal- Notwithstanding its size and robust appearance, B.
low water as well as at depths of 15 to 20 metres. rhoadesii behaves relatively peacefully. Although the
Our B. rhoadesii, which we obtained at a size of males need an area of the aquarium in which they are
approximately 6 to 7 cm, were placed, following a usually found, they do not systematically defend this
short acclimatization period, in a large tank (250 x 70 as a territory. Females are courted but left unharmed;
cm) together with some other haplochromines. First even during spawning other species are tolerated in
we noticed that these juveniles showed a voracious the tank. One thing didn’t, change: their voracious
appetite. After a year of maintenance the fish had appetite; thus we expect further growth of our fish.
reached a size of about 16 cm but did not show any

44
 Corematodus taeniatus Trewavas, 1935
Dr. Andreas Spreinat

A territorial male Corematodus taeniatus photographed near Fort Maguire. Photo by A. Spreinat.

The first Corematodus taeniatus were exported by terest it would slowly approach its prey followed by a
Norman Edwards under the trade name “Haplochromis sudden bite in the caudal fin. Less common were at-
Jacksoni”. In 1988 Stuart Grant exported this cichlid, tacks initiated swimming above the victim. The fin
which was collected at Namalenje Island, as itself was rarely damaged but the fine scales that cover
“Haplochromis Space Mouth”. In November that year most of the haplochromine’s fins made up for a seem-
I received a call from an importer in Bayern that sev- ingly nutritious meal. After a few attacks, however,
eral of these cichlids had arrived, which had immedi- the fin would show rashes and after a few days all
ately aroused my interest. One week after the call I other inhabitants of the tank had scrubbed fins. Mean-
received only a single specimen of the ’hew” cichlid while, the scale-eater showed a healthy appetite for
which had a length of 11 cm. It had a silvery colora- the regular aquarium fare as well.
tion and a partially interrupted, diagonal line. It had a After a while the other fish gradually learned how
very wide mouth in which many rows of teeth were to escape the attacks and kept a distance to the villain.
arranged to form a rasp. It was beyond doubt that this Especially the mbuna proved to learn quickly. It took
cichlid was a member of the genus Corematodus, to about two months before an acceptable balance was
be precisely C. taeniatus. reached. The fins of the non-mbuna were healed and
The other species which is assigned to this genus is the scale-eater had switched almost entirely to the
C. shiranus, a cichlid with faint vertical barring. Both standard aquarium food.
species are known to employ a rather unusual forag- When I introduced a few new, unexperienced cich-
ing technique; they scrape small scales from the pe- lids in the tank they were immediately attacked by C.
duncles and caudal fins of other fish, mainly cichlids. taeniatus. Since they were the only fish in the tank
After a short acclimatization period I placed the unaware of the scale-eater’s habits they succumbed
scale-eater in a community tank with several mbuna the repeated attacks a day later.
and larger haplochromines. Already a few hours after In principle it is possible to house C. taeniatus to-
its introduction the new inhabitant eyed with great gether with other species. Its best tankmates are mbuna
interest the caudal fins of the other fish. The attacks or other robust species like those of the genus
mostly occurred from behind. With a feigned disin- Nimbochromis.

45
 Aulonocara rostratum Trewavas, 1935
Edwin Reitz

A wild caught male Aulonocara rostratum in breeding coloration. Photos by Edwin Reitz.

Until recently Aulonocara rostratum, which has a lake- regularity from the spawning-site. While doing so it
wide distribution, was offered as A. macrochir in the carries small bits of sand out of its nest each time. In
trade, but it is now clear that both these names apply the long term the spawning-site becomes a shallow, clean
to the same species, i.e. A. macrochir is a synonym of dip in the bottom with a diameter of about 40 cm.
A. rostratum. Until 1990 only males were exported as The pre-spawning activity lasts much longer than
females were difficult to find. Then, at the beginning of in any other known Peacock. With my fish it lasts about
1991, the first few females were exported to Europe. two to three days before the actual spawning takes
After two to three weeks of acclimatisation, A. place. In the meantime the male continuously leads
rostratum, like most Malawian cichlids, eats any type the ripe female to its nest.
of aquarium fare and starts showing its splendid col- The male induces the female to deposit some eggs
oration. Although it can grow to a size of about 20 by dragging its lavishly ornamented anal fin over the
cm, it has a very small mouth. In the wild A. rostratum nest while the pair circle around each other. When the
feeds in the typical Aulonocara fashion: it lowers its female deposits some eggs, up to 10 at a time, the
head to just above the substrate and tries to locate any male is parallel to the female (not in T-position). The
prey moving in the sand. Such behaviour is only ob- female turns around quickly and starts picking up the
served in some specimens in the first few weeks after eggs. Meanwhile the male has moved near the eggs
importation. and may fertilize them while they are being picked
A. rostratum lives over the sandy substrates of the up. The male makes another pass and again drags its
lake where adult males dig their nests. A breeding anal fin over the nest, By now the female has col-
colony of this cichlid contains several males and all lected all of the eggs and picks at the spots on the
of them will have constructed a deep crater-nest which male’s anal fin. Then the cycle is repeated. A spawn-
is defended energetically. Therefore one should give ing lasts about one hour.
A. rostratum a large tank with plenty of sand on the During the incubation the female does not eat. Af-
bottom. Interestingly, in the aquarium A. rostratum ter three weeks she releases her fry; in small females
does not seem to construct a crater-nest. I have never they number about 50 but in large, well-fed females
seen it digging. What it does is to take away any ir- spawns of about 150 fry are common.

46
The male drags its ornate anal fin over the spawning-site
and entices the female to lay more eggs.

The female deposits up to 10 eggs at the time. ®

After the eggs are laid, the female turns around and picks them up.

47
 Taeniochromis holotaenia (Regan, 1922)
Peter Baasch

A male Taeniochromis holotaenia in breeding coloration. Photo by Peter Baasch.

Aquarists surely assess a cichlid in a different way to out the years it had never been recognized as an inter-
that used by the experienced ichthyologist, and try to esting cichlid. Now that aquarists are more aware of
“arrange” their fish according to personal points of the complexity of the Malawian species flock a re-
view. The aquarist has “classified” his cichlids accord- newed importation of this species in September 1990
ing their behaviour, gross morphology, and colour was greeted with keen interest.
patterns. Other criteria were not available to the lay- From a morphological viewpoint T. holotaenia be-
man, while scientists applied totally different meth- longs to a group cichlids classified by aquarists as
ods. Both scientist and aquarist had problems appre- “Torpedos”. Recently such cichlids have been exported
ciating each other’s standpoint. The aquarist could not much more frequently than before. Usually they are
understand the reasoning of the scientist, and the sci- put together in an importer’s tank and labeled
entist underestimated the nonscientific approach of the “Haplochromis Torpedo”. Among these “Torpedos”
aquarist. The latter could not understand that one has there are species from different genera, so careful se-
to measure carefully and count certain features before lection from such collections is needed. Cichlids which
a classification of the particular fish could be achieved. are traded as “Torpedos” are e.g. Sciaenochromis gra-
The revision of Malawian haplochromines by David cilis, Sc. spilostichus, Stigmatochromis sp. “Spilo-
Eccles and Ethelwynn Trewavas (1989), and the pub- stichus Type”, Champsochromis spilorhynchus, Ch.
lications of scientists who are aquarists at the same caeruleus, and Maravichromis formosus. All these
time, have resulted in characters like morphology, pig- species are characterized by a very slender body and
mentation pattern, distribution, and evolutionary traits a diagonal line, which in some cases consists of a row
becoming accessible to the hobbyist. The development of smaller spots, on the body. Among this mixture T.
of this mutual understanding may pave the road to a holotaenia is easily identified since it lacks the diago-
better cooperation between the scientist and aquarist. nal stripe.
Taeniochromis holotaenia is one of those species T. holotaenia (holo = complete, taenia = stripe) is
with a unique pigmentation pattern which is crucial characterized by a horizontal band which connects the
for its classification. It is the only species in its genus. eye with the caudal peduncle. In addition to this dis-
Although a few specimens had been exported through- tinct band a black bar runs horizontally between the

48
 T. holotaenia in its natural habitat in Senga Bay (above)
and in the aquarium. Note the difference in coloration.

large aquariums (over 200 cm) and given regular food,

T. holotaenia adapts to its situation and “forgets” about
its piscivorous nature.
eyes, which gives the impression of the fish being Males become territorial simultaneously with the
completely circumscribed by a black line. Depending assumption of breeding coloration. The male con-
on the fish’s, emotions vertical bars may be present as structs a very shallow dip in the sand as spawning-
well. This unique and remarkable pattern formed the site, usually near a rock (sometimes even under one).
basis of placing this species in a genus of its own, The courting process is rather turbulent. Females with
Taeniochromis, although it has some resemblance to a length of about 20 cm release spawns of about 100
species of the genus Dimidiochromis. fry. At release the fry are small but grow quickly show-
T. holotaenia seems to have a wide distribution in ing the characteristic band at an early stage in their
the lake but it also seems to be rare at most locations. development.
The few observations made in its natural habitat were Juveniles have a silvery-yellow ground colour like
restricted to subadults with an average length of about their counterparts in the lake. Adults, however, show
10 cm. They were seen over sand and in the interme- a rather dark pigmentation in the aquarium. Possibly
diate habitat at a depth ranging between 10 and 20 this colour change also occurs in the lake. I shall have
metres. These individuals all showed a very light to wait for the moment the juveniles start changing
ground colour with the distinct black line. Spawning their attire. Male T. holotaenia have a sky-blue breed-
males, or males in breeding colours, were not seen. ing colour with yellowish fins. In this respect it closely
The observations, however, indicated that T. holotaenia resembles Sc. gracilis, Sc. spilostichus and St. sp.
is a pursuit hunter. “Spilostichus Type”.
In captivity this cichlid accepts any regular type of In the aquarium, compared with other cichlids of
aquarium food. The acclimatization of large adults similar length and life-style, T. holotaenia behaves
demands a relatively long period, which seems to be relatively peacefully. Maximum size is around 22 cm
common with piscivores. When kept in sufficiently for males.

49
 Lethrinops micrentodon (Regan, 1922)
Peter Baasch

A wildcaught male Lethrinops micrentodon from Makokola Reef.

Superficially this cichlid seems to belong to the genus A few weeks after I had acclimatized my fish to
Aulonocara rather than to Lethrinops. The latter ge- the confines of the aquarium a male started to stake
nus includes many species which are aquaristically out its territory in one of the corners. The territory
unknown but is getting more and more attention from was demarcated by a wall of sand which had the shape
aquarists. Eccles and Trewavas (1989) revised the of a quarter circle. The radius measured about 45 cm.
genus Lethrinops and described two new genera: If enough room had been given the entire territory
Taeniolethrinops and Tramitichromis. Although there would have been a complete circle with a diameter of
is now a certain logic to the systematics of this large almost one metre! The edge of the construction is just
genus, for the aquarist distinguishing the different spe- a few centimetres high. In the centre of the territory –
cies is still problematic. The females in particular pro- in the case of a quarter circle in the corner – the male
vide a major problem because responsible breeders builds a small cone of very fine sand. This cone serves
want to prevent hybridization by placing the “right” as the spawning-site.
female with a male. It is not known whether L. micrentodon breeds
The species depicted in the photograph above has throughout the year or if it has a specific breeding
been exported only once, in spring 1990, and was iden- season. The specimens in my tank have never stopped
tified as Lethrinops micrentodon. It belongs to a small breeding since I introduced them into the aquarium,
group of deep-living species which have a special- but this behaviour is also known from other Lethrinops
ized feeding behavior. Together with L. microdon and species which breed seasonally in the wild.
L. stridei it feeds predominantly on diatoms which Females, which reach a maximum size of about 10
have settled on the bottom of the lake. Most speci- cm, produce 50 to 60 fry per spawn. The fry are very
mens of these three species have been collected be- small at the time of release but grow rapidly. At a length
low a depth of 50 metres and all three seem to be re- of about 5 cm the males start colouring up. The maxi-
stricted to the southern part of the lake. The L. mum size of the males lies around 13 cm. L. micrentodon
micrentodon that have been exported were collected at is a very attractive cichlid. It requires a small aquarium
Makokola Reef, south of Boadzulu Island. They were (minimum 100 cm) with a sufficient amount of fine sand
found at a depth of 36 m at the border of sand and rock. to provide material for the construction of a nest.

50
 Labeotropheus fuelleborni Ahl, 1927
Peter Baasch

A territorial male Labeotropheus fuelleborni at Katale Island near Chilumba.

Several geographical variants of both species of a yellow, orange or rusty brown coloration on the
Labeotropheus are regularly exported and are among flanks, belly or dorsal part of the body. Some popula-
the popular cichlids from the lake. These two species, tions of L. fuelleborni have a yellow or orange col-
L. fuelleborni and L. trewavasae, live sympatrically oured belly or flank (see photo above of the Katale
at many locations, but hybridisation between them Island population). At one location both species may
hardly ever occurs. I have seen a hybrid between these have an orange colour on the body. In L. trewavasae this
two species in the population at Chidunga Rocks (near colour is restricted to the upper half of the flank whereas
Chipoka) only. As far as I know there are only two in L. fuelleborni it is present on the lower half.
regions where males of both L. fuelleborni and L. These three colour patterns are a part of the genetic
trewavasae are entirely blue. These localities are Mara variation of the two species. The basic colour seems
Rocks and the rocky shores near Nkhata Bay. At all to be entirely blue as this is the colour seen in almost
other locations only one of the two species is present all regions where only one of the two species is present.
or the two species are differently coloured. Most rocky The presence of one of the other two patterns seems
reefs are inhabited by L. trewavasae only whereas at to have originated in a random fashion. It probably
Likoma, Chizumulu, Kande, Mbenji, Namalenje and helped to differentiate between the two species when
Mumbo Island only L. fuelleborni is present. The rocky they came into contact (or it became the only way to
shores along the main coasts harbour both species. differentiate). At each location the process of chance-
There are three major male colour patterns known expression of a pattern and the subsequent sexual se-
for each species. The most common for L. fuelleborni lection developed into today’s checkered distribution
is entirely navy-blue with distinct barring in territo- of the various colour patterns. The colour pattern of
rial males. L. trewavasae is also known in completely L. trewavasae varies more abruptly between adjacent
blue races but the variant at Nkhata Bay is cobalt-blue populations than that of L. fuelleborni. L. fuelleborni
and may thus be distinguished from the navy-blue L. is able to cross sandy areas in shallow water along the
fuelleborni at the same location. The second colour shoreline. L. trewavasae is restricted to the deeper parts
pattern present in both species is a blue body and an of the habitat and may thus never have genetic con-
orange or red dorsal fin. The third pattern consists of tact with nearby populations.

51
 Pseudotropheus saulosi Konings, 1990
Ad Konings

Although large schools of P. saulosi occur, only a small part of the reef is inhabited by this mbuna.

sympatric speciation (the creation of a new species in

the presence of its ancestor), then how do we explain
why that process obviously didn’t occur at Taiwan
Reef. Food is not a limiting factor (at present), be-
cause the reef is inhabited by thousands of rock-dwell-
ing cichlids and is a rich fishing ground for utaka as well.
Pseudotropheus saulosi is one of the commonest
cichlids on the reef and certainly the most conspicu-
ous. Females and non-territorial males forage in large
schools of sometimes more than 100 individuals.
These troups move in the upper reaches of the reef
A territorial male Pseudotropheus saulosi. while picking at the biocover. In November 1990 I
could not find P. saulosi below a depth of 14 metres.
In May 1989, it was the only species around in the
The reef north of Chizumulu Island, the so-called Tai- upper regions of the reef while a very strong current
wan Reef, may prove to be the most important place swept over the reef. Territorial males defend a rela-
in Lake Malawi to study the evolution of the Malawian tively large area in front of a cave, but spawning also
species flock. While most rock-dwelling cichlids have occurred on the open substrate.
formed species complexes at most other locations, P. saulosi shows a strong sexual dichromatism, i.e.
most of these complexes are represented by one spe- males and females have different coloration. The males
cies only at Taiwan Reef. The formation of a species start changing to their blue, zebra-like dress when they
complex elsewhere is probably due to the process of are at a size slightly short of 5 cm (2”). Their maximum
geographical isolation caused by the fluctuating wa- size lies a little over 8.5 cm (3”). In the aquarium, how-
ter level of the lake. Due to its isolated position such ever, they may grow to a size of about 10 cm. Due to its
processes probably did not occur at Taiwan Reef. If small size and vibrant coloration P. saulosi may become
one explains the species complexes by the process of one of most popular cichlids from Lake Malawi.

52
 Labidochromis sp. “Hongi”
Peter Knabe

A mature male Labidochromis sp. “Hongi” claims much of the space in the aquarium when kept with other males.

In 1990 during a collection expedition in Mbamba In the aquarium L. sp. “Hongi” behaves like most
Bay, Tanzania, Hans Fleischer and Thomas Engel dis- Mbuna. The two males and two females which I ob-
covered a beautifully coloured Labidochromis at tained from the first import were placed in a 1000-
Hongi Island which they were able to bring back alive litre aquarium. Within one week the more dominant
to Germany. male spawned with both females. The spawning took
In January 1991 I was able to observe Labido- place on the gravel at a rather randomly chosen site.
chromis sp. “Hongi” in its natural habitat. Hongi Is- The number of fry released after three weeks varied
land, situated between Liuli and Mbamba Bay, con- between seven and 18. The fry grow slowly and reach
sists of two smaller islands which contain mainly very maturity after one year. The parents had a length of
large boulders. These two islets are separated by a 40 about 6 cm when I obtained them; after almost two
m wide and 8 to 12 m deep stretch of water. The biotope years of maintenance in captivity the largest male has
consists here of a sandy bottom with solitary rocks. grown to a length of about eight centimetres. The deco-
Because the two islets consist of large boulders, large ration of the aquarium should have plenty of shelter
caves and overhanging ledges are frequently found for the females.
underwater. In this type of habitat I was able to ob-
serve L. sp. “Hongi” at all levels of the rocky coast.
Most individuals, however, were found at a depth of
about 8 metres. There I saw 8 individuals of which
three were males displaying in front of a cave. As far
as I could observe they fed on plankton or nibbled
form the biocover.
I also found L. sp. “Hongi”, albeit at a lower den-
sity, at Lundo Island. Lundo is, from an aquaristic
viewpoint, a very interesting Island which lies between
Hongi and Mbamba Bay. The shores of Lundo Island
consist of small – maximum football size – rocks. L. sp. “Hongi” in its natural habitat. Photo by Peter Knabe.

53
 VICTORIAN
CICHLIDS

Part I: Introduction to taxonomy and ecology

Ole Seehausen

A male “Haplochromis” nyererei, a zooplanktivorous rock dwelling cichlid.

Although the family Cichlidae is otherwise well-docu- erected genera. Since then about 200 more species
mented in aquaristic literature, Lake Victoria cichlids have been scientifically discovered (most of them still
are one of the few groups which have been given little awaiting description), many of; which bridge the gaps
coverage. between the new genera (Witte, 1974). For these and
For many decades scientific interest in these fishes other reasons (see below) many ichthyologists in the
was also limited (Barel et al., 1991) and only a hand- eighties preferred to refer all species back to
ful of ichthyologists were familiar with them. The tax- Haplochromis as an interim measure.
onomy of Lake Victoria cichlids is still an unresolved Taxonomic classification in haplochromine cichlids
problem. Regan, in his 1922 revision, covered 48 was traditionally based on skull morphology and den-
haplochromine species. Greenwood, in his 1979/80 tition. Striking similarities were found with cichlid
revision, knew about 106, and he redistributed the species from the other African Great Lakes, mainly
former Haplochromis members over 15 new and re- those from Lake Malawi, exemplified by pairs of gen-

54
era like Macropleurodus (Victoria) and Chilotilapia diversity is not reflected in a profound genetic diver-
(Malawi) (Regan, 1922) and Paralabidochromis (Vic- gence. Taxonomy is further complicated by a strong
toria) and Labidochromis (Malawi) (Greenwood, genetically independent variability.
1956a). Such observations resulted in discussions on How much do we think we know about the tax-
polyphyletic versus the previously assumed mono- or onomy and phylogeny (descent) of Victorian
oligo-phyletic nature of the lake flocks (i.e. Green- haplochromines?
wood, 1981). 1. We have strong evidence to assume a mono-
In the last decade molecular biochemical work on phyletic origin for the species flock.
East African cichlids was begun and provided strong 2. Although species differentiation at this evolution-
support for the idea of a monophyletic origin for the ary level is not only difficult in practice, but also a
Victorian haplochromines (Sage et al., 1984; Meyer fundamental problem (Barel et al., 1991), we know of
et al., 1990), with two exceptions being the non-en- many cases in which morphologically and genetically
demic species Pseudocrenilabrus multicolor and almost identical, sympatric, and parasympatric forms,
Astatoreochromis alluaudi. Besides the species of often differing only in male breeding coloration, are
Lake Victoria, the Victorian species flock includes the ecologically distinct species. In several cases it has
haplochromines of Lakes Nabugabo, Kyoga, and very been possible to show their ecological segregation
probably Lakes Edward, George, and Kivu (cf. Meyer (Hoogerhoud et al., 1986; Goldschmidt et al., 1990).
et al., 1990; Greenwood, 1981 for a discussion). This However, this is much more difficult in isolated popu-
flock appears to be the sister group of the Malawian lations.
flock and the two together apparently represent the Why the hesitation to use Greenwood’s new gen-
sister group to Astatoreochromis (Meyer et al., 1990). era? The traditional haplochromine taxonomy is to a
However, the taxonomy at lower levels remains un- large extent based on the anatomy of the feeding ap-
clear because of the extremely low genetic variability paratus. The members of the genus Labrochromis
within the entire Lake Victoria flock (Meyer, pers. Regan, 1920 (re-erected by Greenwood in 1980), for
comm.), The striking feature in the evolution of the example, share a heavy pharyngeal apparatus as the
Victorian haplochromines is that the morphological only derived feature. This was considered indicative

A male “Haplochromis” chilotes, a large species with lobed lips that feeds mainly on insects. Photo by Ole Seehausen.

55
of their monophyletic origin (Greenwood, 1980). Eco- on this basis.
logical fieldwork and experiments in the laboratory I believe that trophic/ecological identification of
have clearly shown that (1) the elements of the feed- Victorian cichlids provides the aquarist with informa-
ing apparatus are phenotypically influenced by envi- tion on his/her fish which is more valuable than that
ronmental factors (eg. type of prey) (Witte, 1984; which can be gained from unreliable taxonomic
Hoogerhoud, 1986), (2) many groups described on the (pseudo)identifications which are often no more than
basis of the anatomy of the feeding apparatus and the the result of guessing. At this point I believe it is nec-
connected skull architecture are in effect ecological essary to stress the fact that identification of Lake Vic-
groups. This does not necessarily deny their validity toria cichlids by coloration alone is impossible unless
as taxonomic units but it is quite possible that parallel one is very familiar with the very species in question
evolution has occurred, producing superficially simi- and even then mistakes are difficult to avoid unless
lar species whose derived characters are convergent other characters are checked.
rather than synapomorphic (features with an identical Lake Victoria is just one of at least six East African
origin which species of the group have in common). cichlid lakes and, being a shallow water body – maxi-
Another reason for the hesitation is that many species mum depth about 90 m – with a huge surface area of
have morphological characters which fall in between approximately 69,000 km² it differs considerably in
those of two or more of Greenwood’s lineages mak- its geography from Lakes Malawi and Tanganyika.
ing it impossible to delimit genera (cf. Hoogerhoud, Comparison of the faunas of the different lakes is
1984; Witte, 1987). an intriguing approach that can provide valuable in-
With the background of the extreme difficulties in formation. However, such investigations have been
identifying and classifying Victorian haplochromines few (Fryer & Iles, 1972; Greenwood, 1981; Witte,
on the one hand, and the increasing need to do so for 1984; Ribbink & Eccles, 1988) because concrete data
adequate fisheries management on the other, research- allowing direct comparisons are still scarce, and re-
ers from Leiden University worked out the concept of cent work on Lake Victoria has shown that statements
trophic groups (Witte & Van Oijen, 1990) which al- based on meagre data have produced an incorrect pic-
lows an ecological classification of haplochromine ture in the past (Fryer & Iles, 1972 versus Witte, 1984
species without any phylogenetic implications. Mem- and Dorit, 1986). Only a few words will be added here
bers of a trophic group are using the same food cat- on aspects of endemism, speciosity, and trophic com-
egory. However, this does not mean that they are feed- position of the flocks.
ing exclusively on the same prey; food choice can dif- Until quite recently it was believed that Lake Vic-
fer seasonally as well as during the 24 hours of the toria cichlids, in contrast to those of Lakes Malawi
day. To identify the trophic status of a series of speci- and Tanganyika, exhibit neither geographic restriction
mens usually a combination of ecological and mor- nor geographic variation (Fryer & Iles, 1972; Green-
phological information has to be considered. wood, 1974). The work of the Haplochromis Ecology
Witte and Van Oijen (1990) further pointed out that Survey Team (HEST) in the south of the lake has
morphological characters correlate more closely with shown that this hypothesis does not hold (Witte, 1984).
the way in which the food is collected than with the Several examples of intraspecific geographic varia-
food type itself. Molluscivores, for instance, which tion were encountered (Dorit, 1986; Witte & Witte-
crush snails between their pharyngeals, are morpho- Maas, 1987) and only a minority of the species has a
logically much closer to some insectivores which also lake-wide distribution (Witte, 1984).
need a strong pharyngeal apparatus, than to other Fryer & Iles (1972) further emphasized that Victo-
molluscivores which pull snails out of their shells with rian cichlids were ecologically less restricted and spe-
their oral teeth. In such cases Witte and Van Oijen cialized when compared to those of the other two lakes.
departed from the strict trophic classification and rec- Recent research proves otherwise. Pronounced eco-
ognized subtrophic groups on the basis of feeding tech- logical segregation and habitat restriction was found
niques. However, in many cases the actual manner in in many cases involving the following criteria (after
which the fish obtains its food is unknown, and it is Witte, 1984): bottom type preference; vertical distri-
obvious that the insectivores, for instance, will have bution along the bottom profile; vertical distribution
to be divided into several subtrophic groups once in the water column; qualitative differences in food
enough information is available. It is my intention to composition, including food size partitioning; quanti-
introduce the trophic groups in a series of articles and tative differences in food composition; differences in
to discuss parts of the Victorian Haplochromis fauna food collection strategy; and partitioning of spawning

56
An orange blotched female of a new species from the “H”. nigricans-group, an epilithic scraper. Photos by Ole Seehausen.

areas. I would like to add behaviour (Seehausen,

1991a).
Even within the geographically very restricted area
of a single bay (Van Oijen, 1981; Barel et al., 1991;
pers. obs.) one encounters completely different spe-
cies assemblages correlated with substrate type and
vegetation. I believe that the comparisons of the spe-
ciosity of the Great Lakes species flocks which have
been published over the years merely reflected the state
of knowledge at the time of publication. According to
Ribbink & Eccles (1988) and Konings (1989) about
“H”. nigricans from an offshore, deep-bodied population,
400 haplochromine cichlids are known from Lake representing the biter type in Haplochromis.
Malawi. Of these about 50% are rock dwelling forms
(Mbuna) with an often very limited geographical dis-
tribution. For a comparison with the Victorian fauna
one has to consider that the existence of a group of
rock-dwelling cichlids similar to Mbuna in Lake Vic-
toria has been known only since 1981 (Van Oijen et
al.). At that time at least 16 rock frequenting taxa were
known from the northern part of the Mwanza Gulf
alone. Sampling was carried out at rock stations by
Sevenster, Bouton, Fermon, and by myself during the
last five years and has revealed many more rock fre-
quenting cichlids within this small region of the lake.
In addition we collected several apparently endemic
“H”. cf. altigenis, one of the larger piscivores which are almost
forms at small offshore islands outside the gulf. extinct, representing the sucker type in Haplochromis.

57
Ribbink and Eccles (1988) found communities of 9 to the enormous impact of the introduction of the nile
14 rock dwelling cichlid species at small isolated rocky perch (Lates sp.) on the indigenous fauna (Barel, 1986;
outcrops in Lake Malawi. This number corresponds Witte et al., 1991). It can be said without doubt that
well with our findings at similar places in Lake Victo- the vast majority of the sublittoral and open water spe-
ria. While being aware that it is risky to draw conclu- cies have declined drastically – about 60% of them
sions from results of geographically restricted work, have not been found for several years – and that the
it appears likely to me that, conservatively calculated, littoral fauna is affected to a lesser degree (Witte et
more than 200 rock dwelling cichlid forms – many al., 1991; Seehausen, in prep.). It is apparent that the
having a very restricted distribution – are to be ex- speed and degree of the decline differed between eco-
pected in Lake Victoria. Ribbink and Eccles (op. cit.) logical groups. Relevant factors are habitat, adult size
reported circa 180 haplochromine cichlid species and abundancy of a species (Witte et al., 1991). Some
found in a Malawian trawl survey of more than 70 details concerning the different cichlid groups will be
stations ranging in depth from 18 to 180 m. Van Oijen given in following articles.
et al. (1981) found circa 200 species in a Victorian
trawl survey of an area covering 45 km2
in the Mwanza Gulf which has a maxi-
mum depth of only 16 m. I can see lit-
tle evidence to assume that the Victo-
rian fauna was less speciose than the
currently known fauna of Lake Ma-
lawi. Unfortunately it is no longer pos-
sible to prove this because of the im-
pact of predation by the introduced
Lates sp. on the flock.
Due to the patchy information a
comparison of the cichlid flocks on
trophic level is also difficult. The pic-
ture of the trophic composition of the
Victorian flock has changed consider-
ably since 1922 and this change shows
rather steadily continuing develop-
ments: a relative decrease in large spe-
cies (i.e. piscivores) and a relative in-
crease in small pelagic or semipelagic
species (i.e. planktivores), deep water
species and Aufwuchs feeders. This
probably reflects the sampling of the
species, which is affected by the fish-
ing techniques employed by research-
ers. Taking the expected increase in
rock frequenting cichlids (mainly al-
gae scrapers and insectivores) into ac-
count, the otherwise pronounced dif-
ference between the Victorian and
Malawian flocks would be reduced
considerably. However, the high per-
centage of piscivores in the original
(pre-Lates) community remains an out-
standing feature of the Victorian flock.
Much of the public and scientific
discussion that has arisen about Lake
Victoria during the last years relates to View of a habitat; rocks and papyrus are often adjacent.

58
View of a rocky habitat under water: grazing “H”. sp. “Velvet Black”. An adult Lates sp, caught by bottom trawl in
the Mwanza Gulf. Photos by Ole Seehausen
References
BAREL, C.D.N. (1986) Endemische Cichliden des Viktoriasees vor haplochromine cichlids from the Mwanza Gulf of Lake Victo-
dem Aussterben. DCG-info, 17 (3). ria. Neth. J. Zool. 31; pp 149-174.
BAREL, C.D.N., W. LIGTVOET, T. GOLDSCHMIDT, F. WITTE & P.C. REGAN, C.T. (1922) The cichlid fishes of Lake Victoria. Proc. Zool.
GOUDSWAARD (1991) The haplochromine cichlids in Lake Vic- Soc. Lond., pp 157-191.
toria: an assessment of biological and fisheries interests. In: RIBBINK, AJ. & D.H. ECCLES (1988) Fish communities in the East
Keenleyside, M.H.A. (Ed.), Cichlid fishes, behaviour, ecology African Great Lakes. In; C. Léveque, MN. Bruton & G.W.
and evolution. Chapman & Hall, London. Sentongo (Eds.) Biology and ecology of African freshwater
DORIT, R.L. (1986) Molecular and morphological variation in Lake fishes. ORSTOM, Paris.
Victoria haplochromine cichlids (Perciformes: Cichlidae). SAGE, R.D., P.V. LOISELLE, P. BASASIBWAKI & A.C. WILSON (1984).
Ph.D. diss. Harvard, Cambridge, Mass. Molecular versus morphological change among cichlid fishes
FRYER, G. & T.D. ILES (1972) The cichlid fishes of the Great Lakes of Lake Victoria. In: A.A. Echelle & I. Komfield (Eds.) Evolu-
of Africa. Oliver & Boyd, Edinburgh. tion of species flocks. University of Maine at Orono Press.
GOLDSCHMIDT, T., F. WITTE & J. DE VISSER (1990) Ecological seg- SEEHAUSEN, O. (1991a) A comparison of some ethological aspects
regation in zooplanktivorous haplochromine species (Pisces: of reproductive ecology in three zooplanktivorous and one gen-
Cichlidae) from Lake Victoria. Oikos (Copenhagen) 58; pp. eralized insectivorous Haplochromis from Lake Victoria. 7th
343-355. Intl. Ichthyol. Congress (The Hague); Bull. zool. Mus. Univ.
GREENWOOD, P.H. (1965) Environmental effects on the pharyn- Amsterdam, special issue, August 1991.
geal mill of a cichlid fish, Astatoreochromis alluaudi and their S EEHAUSEN , O. (1991b) Die zooplanktivoren Cichliden des
taxonomic implications. Proc. Linn. Soc. Lond. 176; 1-10. Viktoriasees. DATZ, 44; pp 715-721.
GREENWOOD, P.H. (1974) The cichlid fishes of Lake Victoria, East WITTE, F. (1984a) Consistency and functional significance of mor-
Africa: the biology and evolution of a species flock. Bull. Br. phological differences between wild-caught and domestic
Mus. Nat. Hist. (Zool.) Suppl. 6. Haplochromis squamipinnis (Pisces: Cichlidae). Neth. J. Zool.
GREENWOOD, P.H. (1981) The haplochromine fishes of the East 34; pp 596-612.
African lakes. Kraus-Thomson Organization, Munich. (Includes WITTE, F. (1984b) Ecological differentiation in Lake Victoria
all the cited papers in reprint unless given separately) haplochromines: comparison of cichlid species flocks in Afri-
HOOGERHOUD, R J.C. (1984) A taxonomic reconsideration of the can lakes. In; A.A. Echelle & I. Komfield (Eds.) Evolution of
haplochromine genera Gaurochromis Greenwood, 1980 and species flocks. University of Maine at Orono Press.
Labrochromis Regan, 1920 (Pisces; Cichlidae). Neth. J. Zool. W ITTE , F., T. G OLDSCHMIDT , J. W ANINK , M. VAN O IJEN , K.
34; pp 539-565. GOUDSWAARD & W. LIGTVOET (1991) Species extinction and eco-
HOOGERHOUD, R.J.C. (1986) Ecological morphology of some cichlid logical changes in Lake Victoria. Proc. 7th Intl. Ichthyol. Con-
fishes. Ph.D. Thesis, Leiden, Netherlands. gress (The Hague), in press.
HOOGERHOUD, R.J.C., F.WITTE & C.D.N. BAREL (1983) The eco- WITTE, F. & M J.P. VAN OIJEN (1990) Taxonomy, ecology and fish-
logical differentiation of the closely resembling Haplochromis ery of: Lake Victoria haplochromine trophic groups. Zool Verh.
species from Lake Victoria. Neth. J. Zool. 33; pp 283-305. (Natl. Nat. Hist. Mus. Leiden) 262.
MEYER, A., T.D. KOCHER, P. BASASIBWAKI & A.C. WILSON (1990) WITTE, F. & E.L.M. WITTE-MAAS (1987) Implications for taxonomy
Monophyletic origin of Lake Victoria cichlid fishes suggested and functional morphology of intraspecific variation in
by mitochondrial DNA sequences. Nature 347; pp 550-553. haplochromine cichlids of Lake Victoria. In: F. Witte. From
OIJEN, M.J.P. VAN, F. WITTE & E.L.M. WITTE-MAAS (1981) An in- form to fishery Ph.D. Thesis, Leiden, Netherlands.
troduction to ecological and taxonomic investigations on the

59
 WEST AFRICAN
CICHLIDS

Lamprologus sp. “Kinganga”

Frank Warzel

This Lamprologus, for once not one of the many new of fish here in 1973. Three years later both scientists
species from Lake Tanganyika, was collected by Heiko published the results of their expedition and in one of
Bleher during one of his many collecting expeditions the many plates accompanying the publication a small
in Africa. The male shown in the photographs, which cichlid is depicted with the caption Lamprologus
is the only specimen of this species collected thus far, werneri. The type species of L. werneri (Poll, 1959:
was caught in the Lower Zaïre (Congo River) near 108-109, Pl. XIX), however, was collected in the lower
Kinganga about 200 km downstream from the Zaïrean part of Malebo Pool, a very shallow, lake-like widen-
capital, Kinshasa. ing of the lower Zaïre river. Malebo Pool is a frequently
The Lower Zaïre at Kinganga had been the site of visited site where exporters collect their aquarium
an earlier expedition by ichthyologists about twenty fishes. As well as Steatocranus casuarius (Buffalo
years ago. An American expedition led by Tyson Head Cichlid), which is regularly exported in large
Roberts and Donald Stewart collected many species quantities from this area, L. werneri, usually under

The cranial gibbosity of the male Lamprologus sp. “Kinganga” is an interesting feature. Photos by Frank Warzel.

60
Lamprologus sp. “Kinganga”, hopefully not the only importation.

trade names like “Lamprologus Congoensis” or cichlid which accepts regular aquarium fare and tol-
“Lamprologus Congolensis”, has also found its way erates any type of water with no noticeable distress.
into many aquaria. Although this cichlid closely re- Even larger fishes find it difficult to intimidate this
sembles the “new” Lamprologus sp. “Kinganga”, es- small cichlid; typical Lamprologus.
pecially in its cylindrical shape, there are a number of
minor differences which suggest that we are dealing References
with a distinct species and not with a geographical
variant of one species, namely Lamprologus werneri. POLL, M (1959) Recherches sur la faune ichthyo-
The male Lamprologus sp. “Kinganga” that I kept logique de la region du Stanley Pool. Annls. Mus. r,
for several years in one of my aquaria, grew to a length Congo Belge. Ser 8º; Sci. Zool. 71; pp 75-174, pls.
of about 10 cm, which is several centimetres shorter XII-XXVI.
than the maximum length known for L. werneri. The ROBERTS, T.R. & D.J. STEWART (1976) An ecological
pattern of light spots on the dorsal and caudal fins is and systematic survey of fishes in the rapids of the
much brighter in Lamprologus sp. “Kinganga” than lower Zaïre or Congo River. Bull. Mus. Comp. Zool.
in L. werneri. Also the vertical markings on the dorsal 147, N 6; pp 239-317.
part of the body are narrower compared to those in L. WARZEL, F. (1990) Ein neuer Lamprologus aus dem
werneri. The relatively large cranial gibbosity of the unteren Zaire? DATZ (43), 2; pp 74-75.
“new” species and the distinct markings on the scales
may be further, possibly typical, characteristics of this
species. Roberts and Stewart noted in their earlier
quoted publication that they too found differences
between these cichlids, and consequently treated both
forms as possible geographical variants of L. werneri.
As an aquarium inhabitant Lamprologus sp.
’Kinganga” is an undemanding, sometimes territorial

61
 Tilapia tholloni (Sauvage, 1884)
Jan ‘t Hooft

The fact that the African cichlids of the genus Tilapia, sisted of the appearance of a red colour on throat and
and the closely related genera Sarotherodon and belly. Shortly after this colour change one of the four
Oreochromis, are relatively poorly represented in showed a further, this time drastic change in colour
aquaristic literature can be explained by their gener- and behaved territorially. Its territory included more
ally large adult size and often destructive behaviour than half of the tank. A few days later a larger indi-
in the aquarium. Tilapiines, however, are very inter- vidual was allowed in its territory and this specimen
esting and I will describe my experiences with one of also adopted the breeding coloration.
the most beautiful among them. Previous reports (Schuitema, 1963; Nieuwenhuizen,
Tilapia tholloni was described by Sauvage in 1884 1967) describe the construction of large spawning pits
under the name Chromis tholloni, honouring the col- in which eggs are laid, but in my tank the pair spawned
lector Tholon. Boulenger (1899) placed it in the ge- before a pit was dug. One day after the second indi-
nus Tilapia where it remains today. T. tholloni is wide- vidual entered the territory the female had attached a
spread in West Africa and is found in Gabon, Congo large batch of eggs onto a piece of bogwood. The pair
Republic, the southern part of Cameroun, and in a part took turns fanning the eggs and dug a large pit. Four
of Zaïre. It is also found in the brackish waters of days after deposition the eggs hatched and the larvae
coastal lagoons although the water of the rivers and were removed to the pit. The wriggling mass was pro-
streams in which most individuals are found contains tected by both parents.
very few minerals. Obviously the water chemistry In the meantime the remaining two fishes formed a
plays an unimportant role. pair as well and spawned within a few days of the first
T. tholloni belongs to the vegetarians in the family pair. This pair too started digging after the eggs were
Cichlidae. Examinations of stomach contents reveal deposited. The spawns of these young couples num-
predominantly remains of plants supplemented with bered more than 500 fry. For months on end the two
insect-larvae and crustaceans. Two other species in
the genus have a similar vegetarian diet. These are T.
zillii and T. rendalli. These three cichlids have a non-
overlapping distribution in most parts of Africa (Thys
van den Audenaerde, 1963; see map).
The specimens I kept in the aquarium were collected
by friends of mine in the Congo Republic. The juve-
niles, with a length of about three centimetres, were
captured in a swampy area just south of the Alima
River, a tributary of the Zaïre (Congo), not far from
the village of Oyo. The water had a depth of 30-60 cm
and the small tilapias could be collected with a handnet
from among the grassy vegetation. After I had intro-
duced four of them in a metre-long aquarium they
immediately showed their vegetarian nature. Within a
The distribution patterns of T. tholloni (yellow), T. rendalli
few days they had completely eaten a thick layer of
(red) and T. zillii (blue).
duckweed which had covered the water in the tank.
Then they started, selectively, on the other plants in
the tank. Some plants like Anubias, Ceratophyllum,
and Nuphar lutea were left untouched, but soft-leaved
plants were devoured in days. I fed them lettuce leaves
and from time to time mosquito larvae and Mysis.
The juvenile T. tholloni had a silvery coloration with
an occasional horizontal line of small spots. At a few
months old, at a size of about six to seven centimetre,
they changed colour. A brown colour with a greenish
hue covered the once silvery body. They were then
placed in a 200 cm-long aquarium in which they
changed colour again at a size of about 10 cm and an
estimated age of about nine months. The change con- A female T. tholloni fanning her eggs. Photos Jan ’t Hooft.

62
pairs spawned. Eggs were always stuck onto a smooth
vertical substrate and the nursery-pit was always dug
one day after spawning.
The four T. tholloni reached a size of about 18 cm
before I gave them away. The maximum size of this
species may be close to 25 cm. The question is whether
this cichlid, under natural circumstances, deposits eggs
onto a hard, smooth surface or digs a pit in which the
eggs are deposited on the bottom. Maybe the spawn-
ing method is dependent on environmental factors and
both methods are practised.

References A juvenile T. tholloni showing its silvery coloration.

NIEUWENHUIZEN, A. van den (1967) Tilapia tholloni.

Het Aquarium. 38; pp 158-164.
S AUVAGE , H.E. (1884) Note sur les poissons de
Franceville, Haut Ogoaué. Bull. Soc. zool. Fr. 9; pp
193-198.
S CHUITEMA , A.K. (1963) Tilapia tholloni. Het
Aquarium. 34; pp 83-85.
THYS VAN DEN AUDENAERDE, D.F.E. (1963) La distribu-
tion geographique des Tilapia au Congo. Bull. Acad.
r. Sci. Outre-mer. 9; pp 570-605.

An adult T. tholloni with neutral coloration.

A pair of wild caught Tilapia tholloni guarding their fry.

63
 Ctenochromis polli and Thoracochromis demeusii
Martin Geerts

The African cichlid fauna can be split up into two main pect haplochromine cichlids in West Africa. Ct. polli
geographical groups: a West African group which, is only found in the old basin of Malebo Pool (Stanley
aquaristically seen, is dominated by Chromidotilapia- Pool). Earlier authors have regarded their specimens,
like cichlids and a East African group dominated by which were caught in the Pool, as representatives of
haplochromine and lamprologine cichlids. Greenwood Th. fasciatus. The maximum length of Ct. polli males
(1987, in Bull. Brit. Mus. nat. Hist. (Zool) 53 (3): 200) amounts to approximately 10 cm. Females remain a
wrote that the chromidotilapiine cichlids are adapted little smaller. The anal fin of the male shows only a
to habitats with flowing water, whereas the haplo- single egg-spot, which was a reason for certain
chromines are noted for their trophic specialisations. aquarists to experiment with this species in order to
Although chromidotilapiine cichlids are absent from find out the function of the spot. The single egg-spot
East Africa, the geographical separation of the two could easily be removed, but investigations indicated
groups is not absolute. This is clearly shown by the that it played no major role at the fertilisation of the
cichlid fauna of the lower Zaïre. Here several eggs.
haplochromine cichlids have adopted a rheophilic life Thoracochromis demeusii (Boulenger, 1899), the
style. The species to which these cichlids belong were West African Humphead, was introduced only recently
revised in 1964 by D. Thys van den Audenaerde, who into the aquaristic hobby. According to Mayland (1989,
recognized four different species: Haplochromis Das Aquarium 244: 613) the Humphead was collected
fasciatus, H. demeusii, H. bakongo and H. polli. In by Heiko Bleher in the lower Zaïre near a village called
his subsequently contested revision, Greenwood Kiganga. Male Th. demeusii grow to a size of at least
(1979) placed the first three species in Thoracochromis 14 cm (Mayland, Cichliden Afrikas: 17-18). Adult
and the last in Ctenochromis. Two of these species males have a cranial gibbosity which led the British
have been introduced into the hobby. ichthyologist Charles Tate Regan to place this cichlid
Ctenochromis polli (Thys v. d. Audenaerde, 1964) in the genus Cyphotilapia. Since its introduction it has
was first introduced in the early seventies. At that time been spawned in the aquarium. Its breeding behav-
it was known under the name “Pelvicachromis Polli” iour is typical of a haplochromine mouthbrooder.
which suggests that aquarists at that time didn’t ex-

A wild caught male Thoracochromis demeusii from Kinganga, Zaïre.

64
 CENTRAL AMERICAN
CICHLIDS

The “Cichlasoma” labridens-complex

Juan Miguel Artigas Azas

The Pánuco river system, México’s second largest At present three species in the labridens-complex
Atlantic drainage and one of the most beautiful found are scientifically described. “C.” labridens was first
in the country, has been pouring its entire flow for described as Heros labridens by the ichthyologist
thousands of years into the Gulf of México. A beauti- Pellegrin in 1903 from specimens collected by pro-
ful assemblage of cichlids of the Parapetenia section fessor Alfredo Duges of the Mexican University of
of the genus Cichlasoma are endemic to the Pánuco Guanajuato in the Rio Verde near the city of the same
basin. One of the species is of uncertain descent and name. The type locality, however, was stated as
has a restricted range: “Cichlasoma” bartoni from the Huasteca Potosina, the name of a large part of the Rio
Rio Verde valley in the western part of the basin. The Pánuco basin. Pellegrin also added Guanajuato to
other species can be placed into two groups, the first Huasteca Potosina as a type locality, something which
associated with “C.” carpinte and the second with “C. is clearly a mistake (there are no cichlid fishes native
“labridens. to Guanajuato state).

The waterfalls at Tamasopo form a boundary in the distribution of some cichlids. All photos by Juan Miguel Artigas Azas.

65
 In 1983 Jeffrey N. Taylor and Robert Rush Miller the fish starts breeding, which, due to the very stable
described “C.” pantostictum and gave redescriptions environment of the springs, takes place throughout the
of ““C.” steindachneri and “C.” labridens. The year. A peak in breeding activity is noted from De-
redescription of the last species was based on a group cember to March. A canary yellow and velvety black
of labridens forms which were collected at over fifty colour pattern then adorns “C.” labridens and this
different locations in the Pánuco system. Dr. Robert change makes it one of the most delightful sights
Rush Miller (pers. comm.) plans to describe another among the New World cichlids (see photo in Cichlids
species in this group, the one known as the Tamasopo Yearbook, vol. 1: 75).
Labridens. This labridens lives in the isolated Rio Pairs then look for a solid surface on which to lay
Gallinas in the mid-western part of the system. An their eggs. Something that would be an easy task in a
estimated twelve thousand years ago the valley of Rio river, but not in the springs where stones are scarce.
Verde – about 10,000 km2 of dry land full of interest- Frequently stems or leaves of water lillies serve as a
ing endemic flora and fauna and situated 1000 metres spawning substrate. Once a site has been chosen and
above sea level – was the site of a large lake. In the thoroughly cleaned, hundred of large – about three
course of time, either erosion or tectonic activity millimetres long – orange eggs are laid, fertilized, and
caused this lake to drain to the Gulf, leaving behind a fanned for two days. After the eggs have hatched the
large swampy area where primitive man is known to larvae are transported to a large pre-dug pit below the
have hunted mammoths. spawning site. From then on it takes another five days
Over the years the swamp almost dried out, leaving before the larvae become free-swimming fry. Pairs will
small swampy regions scattered over the area. These then protect their fry in a ferocious way. The food, in
small swamps survived thanks to the warm water the form of organic detritus, is provided by the female
springs that feed them. The swamps are isolated from by wagging her entire body in the sediment. This
one other, but contain what used to be the fauna of the causes a cloud of debris in which the fry will greedily
prehistoric lake. Two cichlids, namely “C.” bartoni
and “C.” labridens, share their natural habitat in those 1. Pánuco
2. Rio Verde
springs with several other species of fishes. The latter 3. Rio Santa Maria
4. Media Luna
is the so-called Yellow Labridens. 5. Tamasopo
6. Rio Gallinas
“C.” labridens feeds exclusively on crustaceans and 7. Tampaón
8. Tempoal
snails which it finds in the sand and detritus on the 9. Moctezuma
10. Tamesi
bottom of the springs. It has well-developed, molari- 11. Laguna Puerta
form pharyngeal teeth with which it can crush its prey
without problem.
This cichlid, which has a marked ability to change
colours according to its mood, makes an almost reli-
gious ceremony of feeding time by changing its nor-
mally yellowish coloration to a dark, sometimes vel-
vety black pattern with a sprinkling of blue. But this The distribution of “C.” labridens (2, 3, 4), “C.” steindachneri
habit of changing colours reaches its maximum when (5), “C.” sp. “Labridens Tamasopo” (5, 6), “C.” sp. “Labridens
Blue” (7, 8, 9, 10) and “C.” pantostictum (11).

“C.” labridens from the Rio Verde. A natural hybrid between “C.” carpinte and “C.” labridens.

66
forage. The pair take the fry on foraging trips through although present in all of them, may be considered
the habitat, interrupted every now and then by out- rare.
breaks of potential danger in the form of predators or Medialuna spring, the largest of the Rio Verde val-
a curious human cichlid lover. At such moments the ley swamps, presents additional problems for the
male will either face the intruder or, if it is too big, fishes. Man-made irrigation canals, called “asequias”,
flee away. The female will then call her fry and all have been dug from this spring since the seventeenth
will hide under the leaves of water lillies until the dan- century. A large concrete canal that feeds a network of
ger disappears. At dusk as well shelter is found among smaller ones was built by the government in 1977 to
the lillies. Not until the fry have reached two to three take advantage of the large flow of the spring. The
centimetres in length, do they start making solitary new channel intersects the Rio Verde where a differ-
foraging trips, and gradually those that survive will ent fish fauna thrives. During the rainy season the river
leave their parents. At this point they stay close to the may overflow and mix with the spring water in the
overhanging vegetation along the shores of the chan- canal. In this way fishes from the spring can get into
nels which supplies them with shelter. Juveniles are the river and vice versa. “C.” carpinte has thus colo-
also commonly seen around feeding adults where they nized the canal and, although it does not do very well
may learn the technique of stirring up the sediment in the spring, it has hybridized with the yellow
and, while watching, benefit from uncovered small labridens. Moreover it presents an additional threat to
snails or other invertebrates that the adults won’t con- spring cichlids as juvenile carpinte prey upon fry of
sider. “C.” labridens and “C.” bartoni share the habi- the endemic species.
tat, ignoring each other most of the time. This is per- Sarotherodon species have also been introduced to
haps due to the fact that no direct feeding competition the springs as a food fish. It was feared that they could
exists between them. “C.” bartoni, which has conical pose a threat to the naturally occurring cichlids, but
pharyngeal teeth, feeds upon algae on the surface of with the passage of time I have observed their num-
waterlilly leaves or other smooth surfaces, or on bers decreasing. In some springs they have completely
zooplankton. The ratio between both species, how- vanished.
ever, is overwhelmingly on the side of “C.” bartoni The Rio Verde labridens is very similar to the one
and, in fact, the yellow labridens from the springs, found in the springs. It differs mainly in body mor-

A pair “C.” sp. “Labridens Tamasopo” guarding their fry under the waterfall at Tamasopo.

67
“C.” sp. “Labridens Tamasopo” stirring the sediment. A male “C.” sp. “Labridens Blue” from the Rio Pujal.

“C.” steindachneri is the largest cichlid in the Pánuco. A natural “C.” steindachneri x Tamasopo Labridens hybrid.

phology, having a more rounded profile and a shal- The yellow labridens is distributed over the west-
lower head. It has, however, the same breeding col- ern part of the Rio Verde drainage, in the Verde and
oration. The habitat in the Rio Verde is rather differ- Santa María rivers. This area is situated partly at an
ent from that found in the springs. While in the springs altitude of 1000 m above sea level, upstream of the
the labridens are used to very slow flowing water with confluence of the two rivers. It is also found in the
stable temperatures from around 28° C to 32° C and drainage of this area, the Rio Tampaón, until geo-
good visibility, the river has moderately fast-flowing graphical barriers in the form of waterfalls prevent
water – the labridens’ preferred habitat – and signifi- migration further downstream in the system.
cant fluctuations in water level and temperature (from Downstream in the Tampaón river the 102 m high
18° C to 28° C). The visibility in the river is rarely Tamul waterfall provides an effective way of isolat-
over two metres and food is plentiful. ing a unique fish fauna in the Rio Gallinas. Two cich-
The “C.” labridens in the river feed on invertebrates lids from the labridens-group live in this stream in a
which they collect in a similar way to their counter- peculiar and close relationship. One of them is “C.”
parts in the springs. The breeding season is limited to steindachneri, described by Jordan and Snyder in 1900
the months of March through June. Breeding stops and with a stated type locality of Rio Gallinas near
when heavy rains cause much higher water levels with the town of Rascon; the other is the so called Tamasopo
an accompanying decrease in visibility and a lower labridens, a so far undescribed species.
temperature. “C.” steindachneri is a large, slender cichlid with a
As a spawning site, males of the yellow labridens long snout and large conical teeth. It is a piscivore
dig a cave with a depth and width of about 10 to 15 and frequently hunts, livebearers and tetras. On the
cm at the base of a rock, One of the cave’s walls is other hand, “C.” sp. “Labridens Tamasopo” is some-
cleaned and prepared as a spawning substratum. The what smaller, with molariform pharyngeal teeth and a
rest of the breeding behaviour of the river labridens rounded profile. It feeds on invertebrates and collects
matches that of their counterparts in the springs. them in the same fashion as described for “C.”

68
A female “Cichlasoma” pantostictum in breeding coloration (from Laguna de la Puerta).

A male “C.” pantostictum in normal dress. Laguna de la Puerta, the type locality of “C.” pantostictum.

labridens. The estimated ratio between the two spe- dachneri in the Rio Gallinas. Both species breed from
cies is about 100 Labridens for each “C.” stein- December to June with a peak in activity in March.
dachneri. The Rio Gallinas is characterized by its clear During this period the pairs dig caves between the
and well-oxygenated water (pH 7.6; 100° DH!) and rocks and breed like the yellow “C.” labridens in the
has a temperature ranging between 16° and 28° C. Rio Verde. The only difference is the colour of the
“C.” steindachneri can be found only in the Rio eggs which is yellowish instead of orange. Mated pairs
Agua Buena, between the waterfalls at Tamasopo and of the Tamasopo Labridens have a minimum size of
the town of Tambaca, and in Rio Ojo Frio as far as its about ten centimetres for males and seven for females.
confluence with the Rio Agua Buena, where both riv- In contrast, “C.” steindachneri forms pairs only when
ers merge to form the Rio Gallinas. The Tamasopo the male is larger than twenty and the female larger
Labridens exceeds that range to the headwaters of the than fifteen centimetres.
Rio Tamasopo, above the waterfalls. This may sug- The breeding coloration of the two species also dif-
gest that the labridens arrived before “C.” stein- fers. The Tamasopo Labridens has a black and white

69
chessboard pattern with a white forehead. “C.” habitats. The breeding coloration shows the pattern of
steindachneri hardly changes its coloration. the labridens-complex, but this time is all black with
Interestingly, hybrids of the two cichlids exist. A a white forehead. Its breeding habits are difficult to
pair consisting of a male steindachneri and a female assess because of the murky water in which it lives. I
Tamasopo Labridens was photographed underwater have collected “C.” pantostictum in breeding colours
by Ad Konings in March 1991. This explains the oc- from April to June. This cichlid, which grows to over
currence of numerous fishes that have a shape in be- 25 cm in length, has already been spawned in captiv-
tween the two, and these are only found below the ity by Don Danko (Cleveland, Ohio).
Tamasopo waterfalls where both species live together. An interesting note about “C.” pantostictum is that
“C.” steindachneri is the largest cichlid in the two specimens were collected in the head waters of
Pánuco system, in some cases growing to over 40 cm the Rio Sabinas in the upper part of the Tamesi drain-
in size. In some exceptional cases Tamasopo Labridens age. They seemed to live together with the Blue
males can grow to over 30 cm in length but most large Labridens (Darnell, 1962). Whether they were “C.”
males average around 20 cm. pantostictum or somewhat more spotted specimens of
Downstream in the Tampaón river and in the low- the Blue Labridens remains to be seen. I could not
land drainage of the Pánuco (including the Tamesi find “C.” pantostictum in the Rio Sabinas or in other
river, the Pánuco’s northern branch, and the Mocte- affluents of the Rio Tamesi.
zuma and Tempoal rivers, the southern ones), lives a Of all the species in the labridens-complex, in my
different form of the labridens-complex which is own experience “C.” pantostictum is the most aggres-
known as the Blue Labridens. It is distributed over a sive one. All these species require a tank over five
wide area including some small rivers to the south hundred litres and can be housed with other large
that do not belong to the Pánuco system. “Cichlasoma”. Fishes that are too big to fit their
The behaviour of the Blue Labridens is almost iden- mouths, except in the case of “C.” steindachneri, are
tical to that of the previous two labridens. It prefers normally ignored outside breeding times. The large
fast-flowing water with rock-strewn floors for feed- predator of the group will be pleased with all the small
ing. The breeding coloration consists of a blue pattern fish you can provide. “C.” steindachneri is also the
and a red edge on the dorsal and anal fins. The scale- least aggressive of the group.
less blotch behind the pectoral fin, a common feature The foregoing should have given a picture of the
of all species in this group, is blood-red instead of the current status of the “C.” labridens complex. Many
dark-purple observed in all others. questions remain as to whether these forms are to be
“C.” pantostictum lives in isolated coastal lagoons regarded as different species or as geographical vari-
near the Pánuco mouth. Its habitat is very different ants of just one. Nevertheless, they provide the aquarist
from those in which the other species of the labridens- with interesting fishes with great personality.
group are found. The type specimens were collected
in the coastal lagoons “Laguna de la Puerta” and References
“Laguna del Chairel” (type locality), both located near
the city of Tampico. The lagoons are murky with a DARNELL, R. M. (1962) Fishes of the Rio Tamesí and
very low visibility of usually less than half a metre; related coastal lagoons in east-central Mexico, Publ.
the bottoms are always muddy. In lagoons with clear Inst. Mar. Sci. 8; pp 299-365.
water and sandy bottoms “C.” pantostictum could not MEZO, L.M., C. VELASQUEZ, C. ROJAS & J. RIVERA
be found. The temperature of the water in the lagoons, (1989) Rioverde. Ayuntamiento de la ciudad de
which is slightly salty, varies from around 15° C to Rioverde.
28° C. PELLEGRIN, J. (1903) Description de cichlides nouveaux
“C.” pantostictum – its specific name means “spot- de la collection du Muséum. Bull. Mus. Nat. Paris.
ted all over” – differs from the other labridens forms 9; pp 120-125.
by having its entire body covered with small brown TAYLOR, J J. N. & R.R. MILLER (1983) Cichlid Fishes
spots. Such spots can also be found in the Blue (Genus Cichlasoma) of the Rio Pánuco basin, East-
Labridens, but to a much lesser degree. The shape of ern Mexico, with description of a new species. Occ.
“C.” pantostictum resembles most closely that of the paper Mus. Nat. His. Univ. of Kansas. 104; pp 1-
“C.” labridens of the Rio Verde valley. This may not 24.
surprise us given the similarity of the bottoms from
which they feed and the almost stagnant water of their

70
 “Cichlasoma” minckleyi Kornfield & Taylor, 1983
Ad Konings

”Cichlasoma” minckleyi is endemic to the springs at tention from evolutionary biologists as it is a poly-
Cuatro Cienegas in the northern part of México, in morphic species which could give some clues as to
the state of Coahuila. Cuatro Cienegas lies in the east- how polymorphism could be associated with the for-
ern part of the Chihuahua desert. Due to its geographi- mation of species (Kornfield et al., 1982; Liem &
cal location it was isolated from other water systems Kaufman, 1984). “C.” minckleyi is polymorphic in
until men built canals to irrigate land and thus con- its shape and in the structure of the pharyngeal teeth.
nected the springs with the Rio Salado (Rio Grande One morph is deep-bodied and has very large, molari-
system). form teeth on the pharyngeal bones, whereas another
The water system at Cuatro Cienegas consists of morph has a cylindrical shape and sharp, papilliform
many springs (thermal and cold) that drain into lakes pharyngeal teeth (see accompanying photos of at least
or form small creeks or rivers. The water is very hard, four different morphs). Several investigations (Sage
more than 50° DH, and the temperature in a spring I k. Selander, 1975; Kornfield et al., 1982) revealed that
visited in May 1991 was more than 35° C. Originally other genetic characters, which would indicate that
there were seven independent drainage systems of we are dealing with different species, are not coupled
which most drained into closed lakes (Minckley, 1969). to the different morphs . These investigations also
In 1974 only two drainage systems were not intercon- proved that the different morphs interbreed resulting
nected by canals, all others draining into neighbour- in different morphs in one brood. One can thus con-
ing systems or into the Rio Sa1ado (La Bounty, 1974). clude that, even though the morphs look rather differ-
Moreover an extensive underground water system ent, they still recognize each other as belonging to one
connects several springs with each other and through species. After it became known that the Cuatro
these underground channels fish may have migrated Cienegas cichlids represent just one, polymorphic,
from one system to another (Minckley, 1969). The species and not four different ones (La Bounty, 1974),
basin of Cuatro Cienegas was unchanged for a long “C.” minckleyi was used in hypotheses as to how
period of time, maybe for more than a few million (sympatric) speciation in cichlids could take place in
years. Not only cichlids and other fish are endemic to general. One theory (Liem & Kaufman, 1984) hypoth-
this area but also snails, crustaceans, and reptiles. esises that the different morphs would one day mate
“Cichlasoma” minckleyi has received a lot of at- assortatively and give rise to new species. At present,

One of the many springs near Cuatro Cienegas. All photos of “Cichlasoma” minckleyi in this article were taken here.

71
The elongated morph of “Cichlasoma” minckleyi. This specimen was about 18 cm in size.

however, all morphs recognize each other as one spe- lariform morph indicates that there must have been a
cies and the different morphs seem to feed less spe- strong selection for this specialisation. We can hardly
cifically than one would expect from their differences explain this highly advanced specialisation by the vari-
in pharyngeal morphology. ability of “C.” minckleyi alone. A non-assortative
Liem & Kaufman (1984) found it unlikely that “C”. mating system, still present today, would have aver-
minckleyi originated from hybridisation between two aged out the genetic characters promoted by the envi-
species since the molariform morph is so well devel- ronment.
oped as to point to a specialisation. But why hybridi- None of the reports about the Cuatro Cienegas
sation between two species? Couldn’t we explain it cichlid mentions that, beside the trophic morphs, there
by the assumption that an early population of “C.” are also at least three colour morphs. The most com-
minckleyi became split up over several springs and mon in the spring I visited in May 1991 was light-
adapted itself to the local circumstances? Some of blue, the second most common was yellow, and rarely
these springs could temporarily have almost dried out darker, blue-speckled individuals were observed. The
and could thus have started a specialisation towards yellow and light-blue morphs both had deep-bodied
eating snails, an item which was predominantly and cylindrically shaped individuals. The darker blue
avoided by the ancestral population. Some other individuals were small (approximately 5 to 10 cm)
springs could have had a particularly rich population but seem to have different body shapes as well. Along
of small fishes on which the local cichlids could have with this apparently permanent coloration, many very
specialised. “C.” minckleyi seems to be a very vari- dark coloured individuals were observed. However,
able species; in the almost-dried-out spring the indi- when such a dark individual was frightened it quickly
viduals with strong molariform pharyngeal teeth would changed to its basic color. I have made similar obser-
have an advantage over the generalised morphs, vations for “C.” labridens in Rio Florido near Cuidad
whereas in the fish-rich spring fast swimming indi- Valles. Although no in depth examination was made,
viduals would have an advantage. Both these morphs it seemed that only elongated individuals of “C.”
can eat any type of food, as is proven in the laboratory minckleyi turned black and that they did so when hunt-
and by field studies, but the highly specialised mo- ing (small fish?).

72
A deep-bodied, light-blue morph “C.” minckleyi. The hunting(?) colour is fading in this yellow individual.

A deep-bodied, yellow morph “C.” minkleyi. An intermediate morph “C.”minckleyi.

If we combine the variability in trophic characters LIEM, K. F., & L.S. KAUFMAN (1984) Intraspecific
with the presence of different colour morphs then we macroevolution: functional biology of the polymor-
are likely to conclude that these characters have de- phic cichlid species Cichlasoma minkleyi. in: Evo-
veloped independently in isolated populations. There lution of Fish Species Flocks (eds. AA. Echelle and
they could have evolved into highly advanced forms I. Komfield). University of Maine at Omno Press,
without the averaging effect of interbreeding. Only Orono, Maine, pp. 203-215.
after these isolated or even semi-isolated populations MINCKLEY, W.L. (1969) Environments of the bolson of
had evolved into geographical variants, adapted to their Cuatro Cienegas, Coahuila, Mexico, with special
specific environment, were they united by the build- reference to the aquatic biota. Texas Western Press,
ing of canals connecting the once isolated springs. Univ. Texas, El Paso, Sci. Ser. 2; pp. 1-63.
Speciation had not advanced far enough to abolish SAGE, RD. & R.K. SELANDER (1975) Trophic radiation
species recognition among the different populations through polymorphism in cichlid fishes. Proc. Nat.
and thus interbreeding produced the variable popula- Acad. Sci. USA. Vol. 72, No. 11, pp 4669-4673.
tion as we know it today.

References

KORNFIELD, I. L., D.C. SMITH, P.S. GAGNON & J.N.

TAYLOR (1982) The cichlid fish of Cuatro Cienegas,
Mexico: Direct evidence of conspecificity among
distinct morphs. Evolution 36, pp. 658-664.
LABOUNTY, J. F. (1974) Materials for the revision of
cichlids from northern Mexico and southern Texas,
U.S.A.. Ph.D. thesis. Arizona State Univ.

73
 “Cichlasoma” septemfasciatum Regan, 1908
Willem Heijns

The female of the Topaz Cichlid, “Cichlasoma” septemfasciatum, shows a brighter coloration than the male.

Shortly after the creation of Central America by tec- the Atlantic drainage. Once settled they could not cross
tonic activity the narrow stretch of new land was com- the divide, which acted as a barrier that kept the popu-
pletely devoid of a freshwater fish fauna. But, as might lations on either side isolated. Populations derived
be expected, this situation didn’t continue for long. A from one original species could thus develop inde-
great colonisation spread from the south to occupy pendently, sometimes into new species.
the newly created land. Cichlids probably played an This mechanism has led to the evolution of a number
important role as they were among the fishes able to of sibling species, especially in the southern part of
migrate via the sea. Central America. One example is by “Cichlasoma”
How exactly the colonisation took place remains sajica and “C.” septemfasciatum, where the first is
unknown, but the populations of fish entering the new restricted to the Pacific and the second to the Atlantic
land found themselves confronted with a continental drainage.
divide, and they could colonize either the Pacific or “C.” septemfasciatum is distributed over a much
larger area than “C.” sajica. This has resulted in sev-
eral populations which differ mainly in coloration.
Many colour variants of “C.” septemfasciatum are thus
known. An overview of these races can be found in
“Buntbarsche der neuen Welt: Mittelamerika” by
Werner and Stawikowski.
A new population of this species was discovered in
1986 in the Rio Sixaola, in the south of Costa Rica
near the border with Panama. This race of “C.”
septemfasciatum became known as the Topaz Cichlid.
It is characterized by a vibrant yellow-orange colora-
tion which is also visible in males. Initial reports on
this cichlid mentioned that a formal description was
A male of the Topaz Cichlid. Photo by Willem Heijns. on its way but this has never materialized.

74
 Theraps coeruleus Stawikowski & Werner, 1987
Willem Heijns

The female Theraps coeruleus is characterized by the blue spot in the dorsal fin. Photos by Willem Heijns.

In 1985, Stawikowski and Werner discovered a small flowing water. This also could be the reason why T.
cichlid which they named “Small Blue” (Kleine Blaue) coeruleus was not been discovered earlier. One does
and brought it back alive to Germany. In the same not expect to find cichlids in the middle of fast flow-
year, after the authors had published a report about ing streams, and when one does they prove to be dif-
their new discovery, Seeger and Staeck described ficult to catch.
Theraps rheophilus and remarked that the newly dis- T. coeruleus feeds on insect larvae which are found
covered cichlid of Stawikowski and Werner could be among the pebbles on the bottom of the stream. Such
a variant of their species. The “Small Blue” indeed small stones are turned over and the prey secured by a
resembled the specimens which were depicted in the quick bite, preventing it from being carried away by
description of T. rheophilus. Later it became clear that the stream. This behaviour is also observed in the
Seegers and Staeck had been mistaken and T. aquarium, Apart from the crystal clear water they de-
rheophilus was designated a junior synonym of T. mand, T. coeruleus is an easy to maintain and very
lentiginosus. In 1987 Stawikowski and Werner de- attractive cichlid.
scribed T. coeruleus.
The type locality of T. coeruleus lies 30 km south
of Palenque in the Rio Mizol Há Mexico. The river is
a tributary of the Rio Tulija which, in its turn, is an
affluent of the Rio Grijalva. In the description of T.
coeruleus Stawikowski and Werner provide some char-
acteristics by which the genus Theraps can be distin-
guished from other cichlasomini. This forms a basis
on which cichlids that comply with these characteris-
tics can be placed in the genus. The type species of
the genus is T. irregularis. Other species belonging to
the genus are T. lentiginosus and T. coeruleus.
All three species are rheophilic, i.e. they inhabit fast The blue spot lacks in the male T. coeruleus.

75
 Thorichthys pasionis (Rivas, 1962)
Willem Heijns

A male Thorichthys pasionis. Photo by Willem Heijns.

Since the revision of the genus Cichlasoma by species of the subgroup. The basic coloration of the
Kullander (1983) the nomen Thorichthys has become body is violet-blue; the head and lower part of the body
more and more current, in scientific as well as in show many yellow areas. The most prominent char-
aquaristic literature, for species which formed that acteristic, however, is the colour of the throat; dark
section within the original large genus. The genus bordeaux-red to almost black. The colour of the throat
Thorichthys comprises not more than eight species, in T. meeki is red and in T. affinis is yellow. Speci-
five of which form a natural subgroup. T. aureus is mens of T. pasionis in breeding coloration have a black
the best known of this group. The other three form a throat. Normally the colour on the throat is not vis-
natural group as well, of which the Firemouth Cichlid, ible, but this makes its effect more dramatic when the
T. meeki, enjoys great popularity among aquarists. The fish lowers the buccal cavity in a threatening display.
other two species of this subgroup are T. affinis and T. The sharp contrast the suddenly appearing black throat
pasionis. The two subgroups differ in the shape of the forms with the yellow colour on the head must act as
mouth. The lower jaw of T. pasionis and sister species an effective means to deter intruders.
protrudes slightly further than the upper (prognathous) T. pasionis is found in the Rio Usumacinta drain-
whereas the jaws in the species of the other subgroup age (Rio de la Pasion) and in the Lago Petén Itzá in
are equally long (isognathous). The feeding technique Guatemala and southern part of Mexico. T. pasionis
of both groups may therefore differ as well; the prog- is found in streams as well as in lakes. It forages over
nathous species may scoop their food from the sometimes muddy bottoms searching for small prey
substrate whereas the isognathous cichlids pick their such as insect larvae and crustaceans.
choice out of the sediment. However, nothing factual In contrast to some other species of the genus, T.
is known about their natural feeding behaviour. pasionis is relatively easy to keep in an aquarium. They
T. meeki is the best known species of the genus. T. quickly adjust to the conditions of the aquarium and
pasionis is closely related to this cichlid and the only take any type of food offered. The only requirement
difference, besides its distribution, is its yellow col- they have is for clean, oxygen-rich water.
oration. This has led to its trade name of Yellow Meeki.
Indeed, T. pasionis closely resembles the two other

76
 SOUTH AMERICAN
CICHLIDS

Crenicichla species from the Rio Xingú

Frank Warzel

For many, the Rio Xingú is a shallow 2000 km- ready been imported for aquaristic purposes, are
long tributary of the Amazonas which is, by virtue absent from museum collections. This will certainly
of its many rapids, inaccessible to larger boats. For hamper their description in the near future.
ichthyologists and interested aquarists the Rio In the group of nameless cichlids we find sev-
Xingú is the supplier of an inexhaustible array of eral Crenicichla species of which the first was col-
unusual tetras, catfish, and cichlids. Recent inves- lected in 1988 near Altamira by Schliewen,
tigations have indicated that most fishes in the Rio Stawikowski, and Kilian. A few weeks after its dis-
Xingú system are endemic to these waters. A large covery juveniles of this cichlid, introduced as
part of the fauna still awaits a scientific descrip- Crenicichla sp. “Xingú”, were exported from
tion although publications concerning the region’s Belém.
fish-fauna have increased recently. On the other At the same location a year earlier, in the þ sum-
hand it seems that many species, which have al- mer of 1987, Bergleiter hooked two large Creni-

A female Crenicichla sp. “Xingú III” with typical coloration (TL approx. 20 cm). Photos by Frank Warzel.

77
 cichla species that appeared to be unknown to sci-
ence. Reports that a pitch-black Crenicichla was
observed near rocks and in large caves (Schliewen,
pers. comm.) indicated that at least two species were
present. Further imports from Altamira proved that
not one but two other species were to be found there.
These latter two were named C. sp. “Xingú II” and
C. sp. “Xingú III” (Warzel, 1990a).
A fourth species from Altamira was exported in
September 1990 and introduced as C. sp. “Xingú
IV” (Warzel, 1990b). In the meantime this species
has been scientifically described as C. percna
Kullander, 1991. C. phaiospilus is known to occur
in the Rio Xingú but has not yet been exported alive.
1. Altamira; 2. Cachoeira do Espelho, type locality of C. A sixth species was discovered by Harald Schultz
percna; 3. Rio Fresco, type locality of C. phaiospilus; 4. in the Suiá Missú, a tributary of the Rio Xingú.
Cachoeira von Martius, collecting site of C. phaiospilus; S. It is likely that most (if not all) of these six
Suiá Missú, collection site of C. sp, “Suiá Missú”. Crenicichla species are endemic to the Xingú sys-
tem. One of the indicators of their supposed ende-
mism is their relatively high developed specializa-
tion. C. sp. “Xingú I”, for instance, was observed
only in shallow, fast-flowing sections of the river.
It has a very slender body and in the aquarium it
swims in a peculiar oblique position.
In contrast, C, sp. “Xingú III” has a, for
Crenicichla, compressed body with a relatively
large head and eyes. Observations of this species
in the wild and also in part in the aquarium indi-
cate that it prefers caves or dark recesses. C. percna,
Crenicichla sp. “Xingu I”, juveniles (TL c. 15 cm). a species with an extremely shallow head and snout,
is also highly specialized. Although reports about
observations in its natural environment have not
been published, aquarium observations suggest that
we are dealing with a bottom-oriented ambush
hunter. I have observed that C. percna sometimes
rests itself on a rock or on the bottom of the
aquarium, almost feigning to be dead. The usually
light-brown spots on its body then become very dark
and form a conspicuous pattern. Such interesting
behaviour was frequently practised when smaller
Crenicichla sp. “Xingu I”, adult male. fishes could be seen by C. percna.
The specialization, if any, of C. sp. “Xingú II”
and C. phaiospilus is at present unknown. Both
species have a shallow pointed head and high-po-
sitioned eyes. C. sp. “Suiá Missú” from the upper
Xingú has a “normal” shape and is the only species
of the six to have a head profile which resembles
those of C. johanna, C. strigata, and C. lenticulata.
Adults of latter species have a rather rounded snout.
In contrast to the other five species, C. sp. “Suiá
Missú” was not found in the vicinity of rapids or
Crenicichla sp. “Xingu I”, female in breeding coloration. fast flowing water. Lowe-McConnell (1991) found

78
this species in a small lake-like widening of a stream
near Córrego do Gato.
C. sp. “Suiá Missú” has a characteristic colour
pattern which is different from all other Creni-
cichla. The ground colour is grey-green with
round, black spots in the dorsal portion of
the body. These spots are bigger and
merge together towards the end of the
body. Only males seem to have spotted
dorsal fins. The red spots in the spinous part of
the dorsal fin are unique in Crenicichla.
C. percna and C. phaiospilus also show a pecu- Crenicichla sp “Xingu I”, adult female.
liar colour pattern. C. percna (Gr. perknos = dark
spotted) has a leopard pattern with three or four
black spots on its flanks whereas C. phaiospilus
(Gr. phaios = dark, spilus = spot) shows four to five
black blotches on the side.
The pitch-black coloration of C. sp. “Xingú
III” is also uncommon. Only one other
species in this genus is known to have
such a coloration, namely C. cametana
Steindachner, 1911 from the Araguaia
and the Rio Tocantíns which are not far
from the Rio Xingú. This does not automatically
mean that both species are closely related. C. Crenicichla sp “Xingu II”, adult female.
percna, C. phaiospilus, and C. sp. “Suiá Missú” lack
the normally present light-bordered spots at the base
of the caudal peduncle. The colour patterns of C.
sp. “Xingú I” and C. sp. “Xingú II” fall in the nor-
mal range of Crenicichla, although both have ex-
ceptionally intensive red hues, especially in the
unpaired fins. Rarely, and then only in excited emo-
tional phases, a zebra-like colour pattern is seen
on C. sp. “Xingú I” consisting of about 9
to 10 thin, vertical bars which reach half-
way to the ventral region of the body. Out
of all the Xingú species, it is mainly juve-
niles of the latter species that have been imported.
They have an intense orange-yellow ground colour Crenicichla sp “Xingu III”, adult female.
with a pattern of horizontal stripes which remain
to a size of about 20 cm. Such a juvenile pattern is
also known for other species in this group. As long
as these cichlids have this pattern, their behav-
iour is rather peaceful and they
show a tendency to form
schools. It is somewhat more
difficult to mix C. sp. “Xingú
II” and C. sp. “Xingú III” with
other fish. In particular juveniles of C.
sp. “Xingú III” are pugnacious towards conspecifics
and become solitary at an early age. Specimens of
C. sp. “Xingú II” first start to change their colour Crenicichla percna, adult. Photos by Frank Warzel.

79
 pattern at a size of 10 cm. C. sp. “Xingú III” juve-
niles, however, at this size have a completely dark
pattern and are indistinguishable from large adults.
Even the typical iris-crescent behind the pupil is
obvious in small specimens.
Although it may seem like that these six species
have a rather distinct appearance and behaviour,
they all have morphometric characters in common
which place them in a group apart from other
Crenicichla. If one is considering maintaining one
or several of these pike cichlids in the aquarium,
attention should be paid to the individual sizes of
Crenicichla sp. “Xingú II”, juvenile (c. 12 cm TL). these fish. Most grow to a length of more than 30
cm, with the possible exception of C. percna, and
therefore need a large, at least 200 cm long,
aquarium. In addition, these pikes grow enormously
fast and juveniles of 10 cm may add 15 cm in the
first year after importation. Large tanks with plenty
of swimming space are a must, especially for C.
sp. “Xingú I”. The chemical composition of the
water seems to be of minor importance in so far as
the well-being of these cichlids is concerned. Ow-
ing to the fast turnover of food, frequent water
changes are necessary and should keep the buildup
of nitrates to a minimum.
Crenicichla sp. “Xingu II”, female in breeding coloration. Until now none of the Xingú species has been

Crenicichla phaiospilus (above) and Crenicichla sp. “Suiá Missú” (below). Drawings by Frank Warzel.

80
spawned in the aquarium. An analysis of the water LOWE-MCCONNELL, R.H. (1991) Natural history of
near Altamira (Schliewen & Stawikowski, 1989) fishes in Araguaia and Xingú Amazonian tributar-
showed (September, 1988) a temperature of 32.2° ies, Serra do Roncador, Mato Grosso, Brazil.
C, a pH of 6.5, a hardness of circa 1° DH and a Ichthyol. Explor. Freshwaters. Vol. 2 (1), pp 63-82.
conductivity of 120 µS/cm. Even under perfectly SCHLIEWEN, U & R. STAWIKOWSKI (1989) Teleocichla.
imitated water conditions, a spawning could not be DATZ (42) 8; pp 227-231.
induced. Under these circumstances, however, the WARZEL, F. (1989) Neu importiert: Crenicichla aus
females showed their breeding coloration and had Brasilien. DATZ (42) 8; pp 456-457
ripe ovaries. Females of C. sp. “Xingú I” attracted WARZEL, F. (1990a) Neu importiert: Crenicichla aus
males by approaching them in a bent, U-shaped dem nördlichen Brasilien (II). DATZ (43) 12; pp
position of the body, with the central part of the 713-714.
body became very light and conspicuous at the same WARZEL, F. (1990b) Neu importiert: Crenicichla spec.
time. It is known from other Crenicichla that fe- “Xingú IV”. DATZ (44) 1; p 7.
males bend their bodies in an S-shape during the
courtship ritual. Hopefully more specimens will be
exported in near future, so that we may increase
our knowledge of the behaviour and distribution of
these impressive and beautiful pike cichlids.

References

KULLANDER, S.O. (1991) Crenicichla phaiospilus and

C. percna, two new species of pike cichlids
(Teleostei: Cichlidae) from the Rio Xingú, Brazil.
Ichthyol. Explor. Freshwaters. Vol. 1, (4), pp 351-
360.

Crenicichla percna, an adult specimen of unknown sex from Altamira. Photos by Frank Warzel.

81
 Crenicichla sp. cf. regani
Frank Warzel

A female Crenicichla sp. cf, regani from the Rio Xingu near Altamira, Brazil, in breeding coloration. Photos by Frank Warzel.

blance to C. regani. Underwater photographs, taken

by Bergleiter (pers. comm.) in the lower Rio Xingu
near Souzel, showed C. regani in its natural habitat.
The dwarf Pike depicted in the photographs,
Crenicichla sp. cf. regani, was collected by Rainer
Harnoss in a small ditch near Altamira, Rio Xingu
drainage, Brazil, about 200 km upstream from Souzel.
This cichlid differs from C. regani in having a smaller
adult size, a shorter snout, a deeper body, and con-
spicuous markings in the tail. The maximum size of
C. sp. cf. regani seems to be about 6 cm for males and
A male Crenicichla sp. cf. regani.
45 cm for females.
At a length of 4 cm the female assumed breeding
Before I report on Crenicichla sp. cf. regani, another, coloration. Instead of courting her “own” partner she
closely related, dwarf Pike Cichlid – C. regani – should showed off, with her body bent in the typical S-posi-
be mentioned. This attractive small Crenicichla was tion and fins erect, to a much larger male C. regani in
exported in the early seventies and became known an adjacent aquarium. Unfortunately the female ig-
under trade names like “Nanus” or “Dorsocellata”. nored her partner during the entire period during which
Crenicichla regani was described by Ploeg (1989) it seemed that she was willing to spawn. After about
from a large collection of specimens. This dwarf Pike six weeks she lost her beautiful coloration, which is
Cichlid seems to have a wide distribution in Amazonia. common for Crenicichla. Unusually the conspicuous
Ploeg presumed that C. regani was not present in the black and white marking in the dorsal fin disappeared
Rio Negro and that here its niche would be occupied as well. This phenomenon is known only from Pike
by C. notophthalmus. Windisch (pers. comm.), how- Cichlids of the C. lugubris complex; they too can
ever, found dwarf Pikes in the Rio Negro (together “switch off” the coloration in the dorsal fin when they
with C. notophthalmus) which showed a close resem- stop breeding.

82
 Acaronia vultuosa Kullander, 1989
Frank Warzel

Acaronia vultuosa from the Rio Atabapo, Colombia. Photo by Frank Warzel.

Bocca de juquiá (giant-mouthed fish); this is the name mainly at the heavily vegetated edge of the lagoon.
Brazilian fishermen gave cichlids of the genus Habitat descriptions in the literature confirm that A.
Acaronia. They gained this name because of their large, vultuosa prefers stagnant or slowly flowing water. Its
oblique mouth with which they are able to suck in preference for this type of water is further indicated
sometimes rather large sized prey at an astonishing by its typical swimming behaviour: it swims in short
speed. bursts, stopping suddenly while scanning the area for
Besides Acaronia nassa, the newly described A. possible prey. Normally, however, A. vultuosa hovers
vultuosa has also been infrequently imported as an among the weeds and ambushes its prey.
aquarium fish. Both species are identified by their Although Acaronia lives in soft, acidic water, it
characteristic markings on the head. A. vultuosa has a accepts regular tap water without any signs of dis-
very conspicuous pattern consisting of several lines. tress. It takes any type of frozen food but ignores flakes
The pattern in A. nassa has broader bands and round or pelleted aquarium fare. A. vultuosa is not a cichlid
spots which in part have a light edge. The two spe- for a regular community aquarium. It can show many
cies, which show a great resemblance towards each defensive and threatening postures but in general it
other, have a mostly allopatric distribution and are behaves a little shyly towards other fishes, even to-
regarded as sibling species. A. nassa is found in the wards cichlids that are noticeably smaller.
Amazonas drainage while A. vultuosa lives in the A. vultuosa remains smaller than A. nassa; the big-
Orinoco system. Only in the upper Rio Negro have gest specimen recorded measured about 18 cm. The
both species been found sympatrically. This is prob- largest specimens we collected in the Rio Atabapo
ably caused by the Casiquiare which forms a connec- were still a few centimetres short of this length. Fur-
tion between the two river systems. ther observations concerning the behaviour of this
In Spring 1991 we found A. vultuosa in the Rio cichlid have not yet been published. One of these days
Atabapo, south of the Casiquiare. Its biotope, how- I will dim the lights of the tank in order to find out the
ever, was not in the river itself but about 100 m from reason for its large eyes.
the bank in a lagoon fed with water from a spring. We
found many A. vultuosa in different stages of growth

83
 Guianacara sp. “Red Cheek”
Ron Bernhard

A pair of Guianacara sp. “Red Cheek” from Venezuela. Photo by Ron Bernhard.

In 1902, the French ichthyologist Pellegrin described The cichlid which is depicted in the photograph
the cichlid Acara geayi in honour of the collector F. above resembles G. owroewefi in many characteris-
Geay, who caught a number of specimens in the Rio tics. The main feature of this species is the black spot
Camopi in French Guiana. Later the species was placed on the anterior part of the dorsal fin and a black verti-
in the genus Aequidens and until 1989 it was known cal stripe which is most intensively coloured on and
as A. geayi (Pellegrin, 1902). just below the upper lateral line.
The popularity of this cichlid among aquarists Seen from a geographical standpoint the “Red
spawned a vast amount of publications which made Cheek Geayi” is unlikely to be conspecific with G.
the name Aequidens geayi well-known among owroewefi, as it is found in Venezuela whereas G.
hobbyists. owroewefi is distributed over central and east Surinam.
The Swedish ichthyologist Sven Kullander is known These two populations are separated by that of G.
for two reasons. Firstly for his meticulous morpho- sphenozona which inhabits the river systems (mainly
logical examinations of South American cichlids and the Corantijn River) in the western part of Surinam.
secondly for his ability to produce an infinite number G. oelemariensis seems to be endemic in the
of descriptions of species and genera. As early as in Oelemari River where it is sympatric with G.
1980 he indicated his doubts about the placement of owroewefi. Without doubt the “Red Cheek Geayi”
geayi in Aequidens and pointed to the relationship this belongs to the subgenus Guianacara and will be re-
cichlid has with those of the genus Acarichthys. Nine ferred to as Guianacara sp. “Red Cheek”. The “Red
years later Kullander not only introduced a new ge- Cheek Geayi” is one of the most beautiful species of
nus but also two subgenera and three new species! this genus. Its maximum length is about 14 cm for
The genus Guianacara and subgenera Guianacara and males and about 10 cm for females. It has been bred
Oelemaria are described in “The Cichlids of Surinam” in captivity where it proved to be a cave spawner. The
coauthored by Han Nijssen. The species G. geayi, G. female may disappear into the cave for the entire pe-
owroewefi and G. sphenozona belong to the subgenus riod during which the eggs develop into free swim-
Guianacara, whereas the subgenus Oelemaria con- ming fry. The pair bond is strong and they continue to
tains only G. oelemariensis. defend their territory when no fry are present.

84
 Krobia species
Ron Bernhard

The red-eyed Krobia species was collected in French Guiana. Photos by Ron Bernhard.

Before Kullander’s revision in 1989 the genus

Aequidens contained many South American cichlids
which could be placed in several groups. Then
Kullander placed the species which were considered
as the guianensis-group in the new genus Krobia. The
best-known species of the former group was known
to aquarists by the name itanyi, but later it turned out
to be guianensis. These two species, now Krobia
guianensis and K. itanyi, differ from each other only
in their breeding pigmentation patterns. The pattern
of K guianensis changes during breeding to vertical
bars on the body whereas that of K. itanyi remains a
Krobia sp. “Green”, a female fanning her eggs.
horizontal stripe.
When Kullander (1989) defined the new genus he
mentioned that besides the two described species in
Krobia there were two others known to occur in French
Guiana. In the aquarium hobby two undescribed spe-
cies of this genus are known, of which one, Krobia
sp. “Red Eye” was collected in French Guiana. The
other species, K. sp. “Green” came from an unknown
location and has already disappeared from the
aquaristic scene. Krobia species are open substrate
spawners. The eggs are always laid on the same spot
in the territory, which is defended by the pair, even
when no fry are present. The pair bond is strong and
A male Krobia guianensis. Photo by Ad Konings. remains for years.

85
 The “Aufwuchs-feeder”
Roger Häggström

When it comes to the mbuna of Lake Malawi, it is should be a little stiff for easy maintenance. The mesh
well known that they spend most of their time feeding should not be more than 2 mm. Brass or stainless steel
from the Aufwuchs on the surface of rocks. In a nor- screens work as well Cut four bars out of a sheet of
mal community aquarium, though, the fish don’t spend PVC with the sizes 20 x 1 cm and about 2 mm thick.
much time feeding this way. They don’t spend much Glue (PVC glue) the plastic net between two bars on
time feeding in any way. Ever since I have been keep- each side of the long end (see photo). These bars will
ing mbuna, I have wanted to give them the chance to pull the gauze tight over the half-tube. A hole is drilled
feed in a natural way. The result of my deliberations in each end of both plastic (double) bars. The four
is the “Aufwuchs-feeder” which I describe below. screws are mounted in these holes (see photo). On one
This simple device consists of a PVC tube, a square side a rubber band is tied to each screw.
of plastic gauze, rubber bands, four plastic bars, and Now you can load the feeder. Use whatever food is
four rust-free screws. The idea is to put the food be- available as long as it is not too hard or soluble in the
tween the tube (rock equivalent) and the gauze so that water. Pelleted food or frozen food is placed on the
the fish have to scrape the food through the gauze in net and rolled flat using the half-tube. Then take the
order to eat. loose ends of the rubber bands, draw them round the
A PVC tube (20 cm long and with a diameter of 10 tube and mount them on the screws on the other end
cm) is cut lengthwise into two halves which can be of the net. Rinse the whole feeder in running water
used for two “Aufwuchs-feeders”. Drill a hole in one and put it in the aquarium. At first it will take some
end (the “top”) of the half-tube and tie a plastic string time before the fish discover the food, but then you
in this hole. The string should be as long as your tank’s can watch them feeding in their natural way. This may
height plus 10 cm. Tie a handle of plastic or wood at promote the growth of the lips of cichlids like
the other end of the string. This handle should hang Placidochromis milomo or Protomelas ornatus. Now
over the rim of the aquarium thus making it easy to it takes much longer before all food has been eaten. I
remove the feeder from the tank. think this device could be regarded as the equivalent
The plastic gauze (18 x 15 cm) can be purchased in of the wheel in a hamster cage.
some hardware stores. It is used as insect-screen. It

The “Aufwuchs-feeder” let the cichlids eat in a natural way and prolongs the feeding period.

86
 Breeding Tropheus the natural way
Gerard Tijsseling

Tropheus moorii is one of the favourite cichlids from leased in the extreme shallows. These areas have many
Lake Tanganyika. It has been bred in captivity by thou- advantages; they are inaccessible to larger fish (preda-
sands of aquarists. Some of them practise early re- tors) and have plenty of food. Algae-eaters find most
moval of larvae from the female’s mouth in order to of their food in the shallows as these are the lightest
have a higher survival rate of the entire spawn. areas. Small rocks and pebbles provide a substrate for
Among organisms like birds and animals, nature the algae and the necessary shelter.
has provided a process whereby the newly-born indi- The photograph shows how a “shallows” can be
vidual learns some important “tricks” from its mother built onto your tank. Among my friends this setup is
which help it to survive, and which can play an im- known as “tank with balcony”. The platform is glued
portant role at a later stage of its life, e.g. when choos- to one of the tank’s sides. The “balcony” in the photo-
ing a partner. This process is called imprinting and graph has a size of 70 x 50 cm. In my experience 15
usually takes place a few hours or days after birth. cm is the maximum depth for such a platform because
In my opinion fry of T. moorii and similar mouth- deeper “balconies” will be occupied by territorial
brooders from Lake Tanganyika are also imprinted males.
during the first few hours after they have been released The “balcony” is filled with small stones. Care must
from the female’s mouth. In my experience, young be taken to provide oxygen-rich water (place water
which had been taken out of the mouth at a premature inlet over the platform) and lots of light (to stimulate
stage grew up to be bad mouthbrooding females that algae growth). The best thing is: it works perfectly!
swallowed or neglected the first three to five spawns. When a female is about to release her fry she swims
Young which had been released naturally, grew up to up to the “balcony” and stays a week or more with her
be excellent females that brooded the first spawn to offspring before she returns to the group in the rest of
maturity. the tank, The young, however, stay on the “balcony”
In a community aquarium it is hardly possible to till they have reached a size of about two centimetres.
let females release their babies naturally; in most cases Not only Tropheus but also Goby cichlids (Eretmodini)
the fry will be eaten in minutes. Therefore I thought use the “balcony” with success.
of imitating the situation in the lake where fry are re-

The platform is glued to one of the tank’s sides. Photo by Gerard Tijsseling.

87
 The cichlid bible
Martin Geerts

About twenty years ago C. M. Yonge (in his foreword different sources of food.
to Fryer & Iles, The cichlid fishes of the Great Lakes Cichlids have a well adapted feeding apparatus and
of Africa, 1972) described the Great Lakes in East this aspect is discussed by Karl Liem in a separate
Africa as “a field laboratory of evolution”. Although chapter. Liem’s contribution describes the functional
Yonge is a malacologist (mollusc-expert) and Lake morphology of cichlids. Trophic radiation is com-
Tanganyika harbours a great variety of snails, we may monly regarded as the main cause of success in cich-
assume that he was thinking of the cichlid species lids, which is clearly why Liem puts emphasis on the
flocks when he put these words on paper. The East technique with which cichlids collect and process food.
African cichlids have since then been in the limelight His final words, however, show the great need of ad-
of aquarists, ichthyologists and evolutionary biologists. ditional research: “In short, comparative functional
A few years later Dr. P. H. Greenwood (1974, Cichlid morphology has demonstrated the shortcomings of our
fishes of Lake Victoria, East Africa) remarked that the current and generally favoured interpretations of
increased attention given to these cichlids had nega- cichlid diversifications, as monophyletic in origin and
tively influenced the studies of other African fresh- driven by competition in faunistically closed basins.”
water fishes and even of neotropical cichlids. In general biology the species concept is still a hot
The present situation does not seem to be much dif- point of discussion. Greenwood’s chapter concerning
ferent from then as we read in “Cichlid Fishes; Be- this issue is therefore of great interest. With regard to
haviour, Ecology and Evolution”. This book, “the recent investigations on “Cichlasoma” managuense,
cichlid bible”, was published recently by Chapman & he points to the necessity of making a distinction be-
Hall (London). The text is a compilation of papers by tween ecophenotypic variation and polymorphism. On
almost twenty authors, all authorities in their field, page 93 Greenwood expresses his disbelief that the
edited by Miles Keenleyside. The result of their ef- presence of predators, like the Nile perch or the
forts is a must for every “cichlidologist” as it contains tigerfish, would hamper speciation. Even the catas-
all the knowledge concerning cichlids gathered in the trophe in Lake Victoria, caused by the introduction of
last decades. Curiously, this knowledge is not reflected the Nile perch, could not change his opinion.
in handbooks about evolution. The impact the Nile perch had on the species flock
The first chapter of the “bible” treats the intra- in Lake Victoria is further discussed by Barel, Ligtvoet,
familial relationships of cichlids and is written by Goldschmidt, Witte, and Goudswaard. The authors do
Melanie Stiassny. Interestingly, given Greenwood’s not restrict themselves to a description of this eco-
remarks, Stiassny places the cichlids of the East Afri- logical disaster but give information about species
can lakes in a group called “The Rest”. The lampro- identification, ecology, and morphology as well. From
logine cichlids are an exception but during a congress a taxonomic point of view the cichlids from Lake Vic-
held in The Hague (28-8-1991) she expressed her toria seem to provide a so far unsolvable problem, even
doubts as to the correct place this group has in her at the generic level. This situation is also reflected in
cladogram. the treatment of the genus Haplochromis in the re-
That not everything in the “field laboratory of evo- cently published CheckList of the Freshwater Fishes
lution” has its proper place may be illustrated by the of Africa (CLOFFA) (part IV). In this checklist Green-
fact that Ribbink (Chapter 2: 38) regards the mbuna wood’s revisions of the genus are not accepted. Ac-
as a monophyletic group, while Kornfield (Chapter 5: cording to Barel et al. species identification is a fun-
121) speaks of “....as currently defined, the Mbuna is damental problem for which modern taxonomic pro-
a paraphyletic group”. Tony Ribbink treats the ecol- cedures offer no solution (p. 268). The authors find
ogy of the cichlids in Africa’s Great Lakes with au- differences in colour patterns among haplochromines
thority, even though the emphasis is put on Malawian that are morphologically identical. Sexual selection,
cichlids. Yamaoka, in his treatment of the “Feeding therefore, could provide for a necessary reproductive
relationships” (Chapter 7), restores the imbalance by isolation.
referring mainly to Tanganyikan cichlids. Yamaoka’s Mating systems and sexual selection are treated in
contribution is of great interest especially the para- separate chapters. George Barlow discusses mating
graph titled “How to coexist?”. This paragraph also systems among cichlids. He believes that the evolu-
makes clear that much research is still needed to elu- tion of mating systems proceeded from monogamy
cidate the way East African cichlids influence each with biparental care (substrate-guarding species) to
others life styles. Furthermore Yamaoka discusses the polygamy with maternal care (mouthbrooding spe-
techniques with which East African cichlids exploit cies). Barlow comments on many aspects of the dif-

88
ferent breeding techniques and does so with author- cichlids. McKaye questions whether even the nest
ity. Nevertheless a few mistakes have slipped into this building of certain populations of sand-dwelling cich-
chapter. Tilapia rendalli is not a biparental mouth- lids could lead to speciation. He writes “Hence we
brooder as is stated on p. 180. This species is a substrate conclude that bower (= nest. Ed.) form evolves pri-
spawner and that puts the words of Barlow in a differ- marily in response to female choice.” Ad Konings
ent perspective. On the same page Tilapia mariae is (pers. comm.) does not believe so. In his opinion the
treated as a mouthbrooder. Aequidens vittatus (= way a nest is built is mainly dependent on the depth at
Bujurquina vittata) is not found in Surinam. Accord- which it is made. At deeper levels the females see the
ing to Kullander & Nijssen (The Cichlids of Surinam) nests from above which makes the height less impor-
Barlow’s reference relates to Krobia guianensis. tant.
Barlow’s remarks concerning the effects of predation Rosemary Lowe-McConnell describes the distribu-
make it clear that fry and juveniles run the greatest tion and ecology of South American and African
risk of being eaten. Parental care is therefore of ut- riverine cichlids. She gives us an authorative over-
most importance and is treated by Keenleyside (Chap- view of the situation. The last chapter concerns the
ter 9). The author reviews investigations which have breeding of cichlids for food and is written by R. Pullin.
been performed in the field and this makes his contri- “Cichlid Fishes; Behaviour, Ecology and Evolu-
bution of great value to hobbyists. tion”, the bible, is indispensable for every “cichlidolo-
Spawning and guarding of fry demand a coordina- gist”. From the text in the book it is apparent that about
tion between the parents. Mark Nelissen discusses 50% of the species in “the field laboratory of evolu-
communication between cichlids. He distinguishes tion” await formal description, while many genera are
several forms of communication. The importance of in need of a better diagnosis. Moreover nothing is
communication is considered in the chapter written known about the relationships between many differ-
by David Noakes, which deals with the ontogeny of ent groups of species. This may be the reason why
cichlid behavior. Kenneth McKaye comments on cichlids, a prime example of speciation, are so poorly
sexual selection (Chapter 12). Sexual selection is cur- represented in books about evolution.
rently regarded as the prime factor in speciation among

The nest of Cyathopharynx furcifer, here the variant in Moliro Bay, varies in height according to the depth at which it is built.

89
 Cichlid classics
Mary Bailey

Recent years have seen several major revisions of nine, or neuter – which, in classical Latin, is usually
cichlid genera, with, in most cases, the original genus recognisable by the ending of the word. The best
being restricted to a few of its former species, and known are “-us” (masculine), “-a” (feminine), and “-
new genera being erected for the remainder. Frequently um” (neuter). Unfortunately in taxonomy it is not quite
the gender of the new generic names has differed from that simple, as if a generic name is a latinised form of
that of the original genus, necessitating alterations to a word from another language, it retains its original
some of the specific names as well. As Latin is no gender (if any). Thus Cichlasoma, which has Greek
longer generally taught in schools the average aquarist “roots”, appears feminine at first glance because of
has been left in a state of some confusion, and it is its “a” ending, but is in fact neuter. It may be neces-
hoped that this article will cast a little light upon the sary to refer back to the original generic description
subject. in order to establish the derivation, and hence the gen-
The language of taxonomy is nominally Latin, but der, of a generic name. Aquarists who find themselves
many of the word used are latinised forms of words confused may take heart from the fact that even the
from other languages (mainly Greek) and proper greatest taxonomists can themselves sometimes be in
names from all over the world. The grammar of tax- error – the Greek word “gramma” can mean “letter”,
onomy is, however, strictly Latin. Under the trinomial in which case it is neuter, or “line”, when it is femi-
system of nomenclature, devised by Linnaeus in the nine; Dr. C. T. Regan confused the genders of the two
18th century, all living beings are classified by three when erecting Apistogramma, and, while meaning
names (the meaning of trinominal): the generic, the “line”, used the neuter gender. This error remained
specific, and the subspecific. All generic names are uncorrected until Kullander (1980) revised the genus.
nouns (words which name a thing); “fish” is a noun If we wish to refer to a particular member of a ge-
which tells us with what part of the animal kingdom nus then we must use the specific name, eg Tropheus
we are dealing, and likewise Tropheus, Etroplus, duboisi. The specific name “qualifies” the generic;
Aequidens, are Latin nouns, generic names which “duboisi” tells us that we are dealing with a particular
specify particular groups of fish species. type of Tropheus, that named after a M. Dubois. It
Latin nouns all have a gender – masculine, femi- enables us to refer to this species in the literature, and,

The specific name of this cichlid, Ophthalmotilapia nasuta, is an adjective meaning “having a (distinctive) nose” and ends
according to the gender of the generic name, which is feminine. Photo taken in Moliro Bay, Zaïre.

90
to many of us, conjures up an immediate visual image must be altered. Thus Cichlasoma severum (neuter) is
derived from our own data banks. Subspecific names now Heros severus (masculine).
allow us to identify distinct genetic groups within a A genitive is the possessive form of a Latin noun,
species when such exist; they follow the same gram- and is equivalent to “of X” in English. In taxonomy it
matical rules as specific names. The specific name can is almost always used to name a species after a per-
be one of three types: an adjective (a descriptive word, son, although there are rare exceptions – Tanganicodus
eg “auratus”, meaning “golden”); a genitive (the pos- irsacae is named for the organisation IRSAC, and
sessive of a name, eg “duboisi”, which means “of Tangachromis dhanisi after a boat, the “Baron von
Dubois”); or a noun in apposition (a noun used to Dhanis”. Although as a rule adjectives are used to de-
qualify another noun, eg “Mr Bun the Baker”, or, note place names, occasionally the genitive of such a
aquatically, Pseudotropheus zebra, “Cichlasoma” name is used, usually where it is impossible to form a
centrarchus). sensible adjective, as with African names such as
The commonest type of Latin adjective has the “- Nkhata and Nkamba.
us”, “-a”, “-um” endings (respectively masculine, Genitives take the gender of their originator, with
feminine, neuter) mentioned earlier in respect of nouns; the masculine ending “-i” (”regani”, “burtoni”), and
there is another common type with the endings “-is” feminine “-ae” (”kingsleyae”, “trewavasae”). The
(masculine and feminine) and “-e” (neuter), eg feminine ending has also been used in the place name
Apistogramma pertensis, “Cichlasoma” nicara- derivatives “nkatae” and ’hkambae” for reasons of
guense. It is worth noting in passing that in Latin ALL common sense and pronounciation*.
syllables are pronounced, and that the correct Very very occasionally a species is named after two
pronounciation of the “ense” ending is “en-say” (two people, thus we have “Cichlasoma” hogaboomorum,
syllables)! named for the Hogaboom brothers, “-orum” being the
Adjectival specific names MUST agree in gender masculine plural ending. The feminine equivalent is
with the generic name to which they are applied, and “-arum”, but I can think of no example of its use.
so if the species is moved from one genus to another, Genitives are thus quite distinctive, and unlikely to
with an attendant change of gender, the specific name be confused with adjectives. They remain constant

Tanganicodus irsacae is named after an organisation. Photo taken near Kavalla Island, Zaïre.

91
whatever the gender of the generic name. languages. If it is to fulfil this purpose then it needs to
Unfortunately nouns in apposition often have “-us”, be used accurately and precisely. And, in an era where
“-a”, “-um” endings like some adjectives, and when scientists are beginning to take notice of observations
the species involved are moved to a different genus it made by aquarists, it is important for our credibility
is all too common for these names to be subjected to that we too are precise and accurate if we wish to make
’blanket” gender changing by aquarists. But because a meaningful contribution to the knowledge of our
they are nouns in their own right they have their own cichlids.
gender, which always remains the same whatever the
generic name. Some are easy to spot, especially where References
they are the names of other animals – the derivation
of Pseudotropheus zebra is obvious to everyone; if KULLANDER, S.O. (1980) A taxonomical study of the
you know that Centrarchus is the name of another genus Apistogramma Regan, with a revision of Bra-
genus which ’Cichlasoma” centrarchus resembles zilian and Peruvian species (Teleostei: Percoidei:
then there is no problem, But there are also “ordinary” Cichlidae). Bonn. Zool. Monog. 14: 22.
nouns used in apposition, and, to exacerbate matters, UFERMANN, A., R. ALLGAYER, AND M. GEERTS (1987)
some of these are themselves composites of a noun Cichlid Catalogue. Brumath, France.
and adjective. “Melanotheron” (”black-chin”) and
“curviceps” (”curved-head”) will probably be left well * For many years there has been confusion over the
alone, but the same cannot be said for “maculicauda” genitive form of the name Moore (referring to J.E.S.
(”spotted-tail”) or “longimanus” (”long-hand” [-fin]) Moore, collector of the type species of the genus Tro-
which look like adjectives but are not. Life becomes pheus). Boulenger’s original description uses moorii,
even more confusing when one realises that there is a being the genitive of the latinised form “Moorius” of
perfectly good adjective “maculicaudatus, -a, -um” Moore’s name; some authors maintain that “moorei”,
(”having a spotted tail”) which must agree with any a simple genitive of the English form, should be used.
generic name to which it is applied. If, however, we refer to the rules of zoological no-
It must be quite obvious, from the above, that with- menclature we find that there is no room for dispute.
out a lot of background knowledge, either of Latin Any author describing a new species can choose
and Greek vocabulary, or of taxonomic literature, the whether he latinises a name before using it, or simply
average aquarist cannot hope correctly to revise every applies the appropriate ending. It is not for subsequent
specific name where generic change has occurred. In authors to question his decision. The name published
consequence it is wise to conduct a little research be- in the original description of a taxon is the correct one
fore committing oneself to paper or in public. (provided, of course, that there is no question of syn-
Recently some authors have adopted the practice onymy, preoccupation, or gender change). A name can
of explaining the derivations and genders of the names be altered only if it was misspellt by the typist or printer
they create; many earlier authors, however, omit to (but NOT if the mistake was the author’s) or if it is
provide any derivation, and gender/grammar are not grammatically incorrect. None of these apply in
even mentioned as these used to be a matter of com- Boulenger’s case, so his published name Tropheus
mon knowledge amongst scientists, who, until com- moorii is the correct one.
paratively recently, would automatically have received
a “classical education”. In the absence of any clues in
the original description it can be helpful to examine
the subsequent synonymy of a species, as any earlier
variation in the specific name will usually point to an
adjective. The .”Cichlid Catalogue” of Ufermann,
Allgayer, and Geerts (1987) can be extremely helpful
for this purpose. Doubtless many aquarists will ques-
tion whether there is any point in going to all this trou-
ble.
Well, the point of any language, taxonomic or oth-
erwise, is to make oneself understood, and the trino-
mial system is intended to facilitate the communica-
tion of species identity between people of different

92
 Spawning techniques in mouthbrooders
1. Fertilization in mouthbrooding cichlids
Dr. Ethelwynn Trewavas

Wickler (1962a,b) put forward two theories to explain ily Cichlidae, in its most advanced form in the tribes
the evolution of the egg-mimicking spots on the anal fin Tilapiini and Haplochromini (Peters & Berns, 1978 and
of males of many mouth-brooding haplochromines: 1. 1979). The tilapiines include both substrate- and mouth-
that they served to trigger spawning in the female, and 2. brooders. Evolution from the former to the latter involved
that the female, who has taken the eggs into her mouth the regression and loss of features adaptive to substrate-
very rapidly as they are laid, is tricked into trying to pick spawning and guarding, and the development of new
up the egg-dummies and so to inhale sperm from the features appropriate to the care of larger and fewer,
male’s vent and to ensure the oral fertilization of any nonadhesive eggs. The problem was, how to get the eggs
eggs not already fertilized. These theories have been rapidly into the maternal mouth without neglecting their
widely accepted, but recently they have been challenged fertilization. Meeting this need there evolved a behav-
by Eva Hert (1989). Hert’s experiments showed that the ioural trait, first described by Baerends & Baerends-Van
spots in A. elegans attract and stimulate the female, al- Roon (1950: fig. 50, 51) and named “tail-chasing” (Fryer,
though only if accompanied by courting movements of 1959), T-positions (Heinrich, 1967), making half-loops
the male. The fertilization rate, however, was not sig- (Balon, 1977) or “tight circles” (Kapralski, 1990b) with
nificantly different between males with egg-spots and each other. The alternating T-positions are: TA: male’s
those from which the spots had been removed by freeze- snout contacts female’s body near the vent, TB: male
branding, both achieving nearly 100%. places his body before the female with his vent near her
mouth. The series of T-positions culminates after a very
Comparison of experimental with natural conditions short time or up to 30 min. (Berns et al., 1978) in the
“real spawning”, when at TA the female lays an egg or a
The experimental conditions were: one male and batch of eggs and at TB she collects them, either allow-
five females were together isolated in each of 12 sepa- ing the male to fertilize them first or more rapidly so that
rate compartments. There were thus no rival males, she has to inhale the sperm to fertilize them in her mouth.
no predators, no egg-robbers, no water currents. These It is this last movement, oral fertilization, that Wickler
are ideal conditions for undisturbed spawning. In con- suggested was prompted by the deception of the egg-
trast abundant evidence exists that in nature a spawn- mimics on the male’s anal fin in species possessing them.
ing pair may be surrounded by any or all of these haz- The series of T-positions (TA-TB-TA-TB) is repeated
ards (see McKaye, 1983; Ribbink et al. 1983; until the female has no more eggs to lay. The importance
Konings, 1989: 291; 1990: 150, 204). of circling with T-positions is that by mutual stimulation
it synchronises the spawning acts of male and female
Adaptation to mouthbrooding and facilitates fertilization either immediately before,
during or after the collection of the eggs (Mrowka, 1987).
Mouthbrooding has evolved several times in the fam- Wickler’s prime exemplar of oral fertilization was

A pair of Copadichromis eucinostomus (Msuli Point, 16 m) displaying the TB-position; see text for explanation.

93
Astatotilapia (formerly Haplochromis) burtoni. His film semble them in colour and shape (Goldschmidt, 1991),
of this species shows the female making over the “egg- The contrasting border is well-marked and regular (or
dummies” the same lip-movements that, immediately the fin itself is dark). Their mimicry has been accepted
before, she has used to suck in the real eggs. Descrip- even by some who reject Wickler’s explanation of it.
tions of spawning in mbuna by careful aquarists give the Among the mbuna there are some with a pattern indis-
same picture, e.g. Mary Bailey (letter of 28-1-1989) de- tinguishable from this type, others in which the spots are
scribes the apparent efforts of Pseudotropheus socolofi much smaller and arranged in a cluster instead of in rows.
to suck in the eggsspots as “persistent”. Wickler used Occasionally they may occur also on the dorsal fin or in
the word “energisch” for the same behaviour in his “H. females. There is interspecific variation as well as a de-
wingatii” (the species Wickler studied was Astatotilapia gree of uniformity within a species or a population
cf. bloyetii from Tanzania). (Konings, 1990 and pers. comm.). This gives some sup-
The female “H. wingatii”, however, sometimes gives port to the theory of Axelrod (see Ribbink in Jackson &
the male time to fertilize the eggs before she collects Ribbink, 1975) that the female recognizes the species-
them. In that case she “does not deem the spots worth a specific pattern. This theory and that of Konings (1989,
glance” (Wickler, trans. E.T.). This indicates that when p. 29) that the displayed anal fin, especially when bear-
she does take an interest in the fin her behaviour is di- ing a brightly coloured pattern, “shows the female where
rected to fertilization rather than to egg-safety. Wickler’s to spawn” are applicable both to patterns not resembling
theory, on the other hand, distinguishes between the fe- eggs and to egg-spots. Neither claims any influence of
male’s probable motive (safety) and the survival-value the spots on oral fertilization.
of her action (fertilization).
It is evident that “H. wingatii” is flexible in its tech- Oral fertilization without egg-spots
nique, as, in response to environmental situations,
some other species are found to be (Konings, this vol. For haplochromines see Konings, this vol. p. 97. Fer-
p. 97). A. burtoni seems to be more rigid in its use of tilization in maternal mouthbrooding tilapiines has been
oral fertilization. Paulo’s experiment (1975) with this described for seven species among the three subgenera
species, in which he removed a male’s anal fin, clearly of Oreochromis. These accounts are summarized in
shows that the egg spots, and even the fin itself, are Trewavas, 1983 (pp. 190, 367, 387, 419, 450, 475, 487),
not necessary for oral fertilization. This cannot be de- where references to the original records are supplied. In
duced from Hert’s results because we do not know six species oral fertilization is inferred from the snap-
whether A. elegans used oral fertilization either dur- ping actions of the female close to the genital papilla of
ing the experiment or otherwise. the male during the laying and collecting of the eggs.
The use of similar lip-movements over eggs and The seventh species, O. (Nyasalapia) karomo, was ob-
“egg-dummies” which seemed to support Wickler’s served in the Malagarazi Swamps, where male and fe-
theory, is weakened as evidence when we realise that male glided successively over the nest evidently laying
the same method, namely suction, would be used for and fertilizing the eggs. Similar behaviour by O.
taking in either eggs or sperm. At this time the fin is macrochir was seen in the Lufira River, but in the labo-
probably covered with sperm. ratory the female mouthed the genital tassel of the male.
Wickler suggested that the tassel might function as a kind
Occurrence of egg-spots of spermatophore. If so, it would be by retaining the sper-
matic fluid (slightly more viscous than water) at the sur-
Tilapiine mouthbrooders and some haplochromines face of its branches long enough for the female to obtain
have no bright yellow spots on the anal fin of males. a concentration of sperm. The male’s genital pore is at
Many Malawian species have a conspicuous pattern, the base of the bifid tassel (see Trewavas, 1983, fig. 171).
usually yellow, that does not resemble eggs, but in mbuna The tassel in all species of this subgenus (Nyasalapia)
and some others the orange to orange-yellow spots are is cream- to orange-coloured in contrast to the usually
surrounded by a contrasting border or a dark fin and have black breeding colour of the male. The black male of O.
been called egg-spots. The closest resemblance between mossambicus has a prominent yellow papilla. In O.
spots and eggs characterises the haplochromines of the (Alcolapia) alcalica grahami the white papilla is sur-
Lakes Victoria, Edward, and Kivu (VEK) group and re- rounded by a black ring, which is highly conspicuous on
lated species of streams, lagoons and the inshore waters the white skin of the belly as shown in a photograph by
of lakes, e.g. A. burtoni and A. calliptera. In these the Dr. Wickler in Albrecht et al. (1968), also reproduced in
spots are of the same order of size as the eggs and re- Trewavas, 1983. As Fryer & Iles (1972) note, conspicu-

94
 yellow pattern. The female is targeting this fin. The
second is a photo, by Milner, on p. 4 of Jackson &
Ribbink (1975), of a group of partial albino Pseudo-
tropheus zebra. These have no dark pigment – even
the retina shows pink through the pupil – but in three
individuals the anal fin bears a cluster of 2-4 small
orange-yellow spots in a black field. Nothing could
be more conspicuous or less egg-like. What genetic
quirk has here made an exception in favour of the most
valuable asset of a male cichlid?

Summary and conclusions

Astatotilapia calliptera, Thumbi East, L. Malawi. The perfect egg-
mimics. Note their position. See also photos on pp 54-57.
In maternal mouthbrooding cichlids fertilization, es-
pecially oral fertilization, is facilitated by various
means. Mutual circling with T-positions ensures in-
semination either during or after collection of the eggs.
The female is guided to the location of the male’s geni-
tal papilla by a conspicuous device, – the colour of
the papilla in contrast with its surroundings, an out-
growth of the papilla (rosette or tassel) or a conspicu-
ous pattern on the male’s anal fin. It is suggested that
the tassel or fin delays the dispersion of the somewhat
viscous seminal fluid and that it is for this that the
female mouths tassel or fin at the end of a spawning
bout if she still holds unfertilized eggs in her mouth.
Aulonocara sp. “Chitande Type Masinje”. A lavish display
This theory of the female’s interest in the male’s fin
on a black fin.
does not differentiate between the egg-pattern and any
other, and causes us to look again at Wickler’s second
theory. An emergency that causes the female to pick up
the eggs unfertilized may also cause her concern for egg-
safety to amount to an obsession and make her more
easily deceived by the egg-dummies. Some studies of
fertilization already show that behaviour in this function
may be influenced by the environmental situation (see
Kapralski, 1990a and c) and more such studies are desir-
able, using species with different anal-fin patterns. The
egg-pattern appears to be very successful, since it oc-
curs in all species of Astatotilapia and related genera
constituting the VEK group of haplochromines as well
Petrochromis sp., Bulu Point, L. Tanganyika. Egg-spots on as several species of the more diverse flock of Lake
both anal and dorsal fins. Photos by Ad Konings.
Malawi and some of Lake Tanganyika. Wickler’s two
theories are the only ones that offer an explanation of its
ousness is an attribute shared by these features and the evolution and they have so far not been disproved. Un-
decorated anal fin of haplochromines. See also less at least one of them is accepted, then I doubt if we
Goldschmidt (1991), who seems to rate it as at least as are justified in giving the resemblance the name of egg
important as the mimetic nature of the egg-spots. mimicry. After all, it is a very simple shape and the dark
As food for thought on this subject I call attention border that gives the spots a three-dimensional appear-
to two published photos. One, by Spreinat (1990: 529), ance also makes them very conspicuous, a valuable as-
shows a pair of Rhamphochromis sp. in an early stage set. The theory of mimicry must rest on the supposition
of courtship. The body and fins are of uniform colour, that the egg-image has a special significance for the
except the male’s anal fin, which bears a conspicuous gravid female, and this has not been disproved.

95
 Cost-benefit estimate KAPRALSKI, A. (1990c) Breeding Haplochromis compressiceps.
TFH August. pp 32-34, 37, 41.
It has been objected (Konings, 1989: 30) that the KONINGS, A. (1989) Lake Malawi cichlids in their natural
conspicuously decorated anal fin may carry the dis- habitat. Verduijn Cichlids & Lake Fish Movies. Nether-
lands. 313 pp. illustr.
advantage of attracting predators, especially when the
KONINGS, A. (1990) Cichlids and all the other fishes of Lake
spots are egg-mimics. This is a special case of a di- Malawi. TFH Publ. Neptune City, NJ. 495 pp. illustr.
lemma very common throughout the vertebrates. Evo- MCKAYE, K.R. (1983) Ecology and breeding behaviour of a
lution seems to have decided that the benefit of suc- cichlid fish, Cyrtocara eucinostomus, on a large lek in Lake
cessful breeding outweighs the possible cost. This is Malawi, Africa. Env. Biol. Fish. 8; pp 81-98.
why both we and the peahen can enjoy the display of MROWKA, W. (1987) Oral fertilization in a mouth-brooding
the peacock, and why both female cichlids and fish cichlid fish. Ethology. 74; pp 293-296.
photographers are seduced by the brilliant colours of PAULO, J. (1975) Bedeutung der Eiflecke. DCG-Info (Ger-
male cichlids. man Cichl. Ass.) 6; pp 163-165.
PETERS, H.M. & S. BERNS (1978) Über die Vorgeschichte der
maulbrütenden Cichliden. 1. Was uns die Haftorganen der
Acknowledgments
Larven lehren, pp 211-217. 2. Zwei Typen von Maul-
brütern. pp 324331. (illstr. in col.). Aquarium Magazin
I have benefitted by discussions with Dr. Tijs PETERS, H.M. & S. BERNS (1979) Regression und Progression
Goldschmidt and with aquarist friends, especially in der Evolution maulbrütender Cichliden. Mittl. Hamb.
Mary Bailey and Ad Konings, as well as from their zool. Mus. Inst. 76; pp 506-508.
published works. I am grateful to my friend Mrs. Isabel RIBBINK, AJ., B.A. MARSH, A.C. MARSH, A.C. RIBBINK, AND
Rampton for kindly typing the script. B.J. SHARP (1983) A preliminary survey of the cichlid fishes
of rocky habitats in Lake Malawi. S. Afr. J. Zool 18 (3); pp
References 149-310.
SPREINAT, A. (1990) Beobachtungen an Rhamphochromisarten.
ALBRECHT, H., R. APFELBACH & W. WICKLER (1968) Über die DATZ 43; pp. 528-533.
Eigenständigkeit der Art Tilapia grahami Boulenger, ihren TREWAVAS, E. (1983) Tilapiine fishes of the genera Santher-
Grubenbau und die Zucht in reinem Sueßwasser (Pisces, odon, Oreochromis and Danakilia. 583p., London, Br. Mus.
Cichlidae) Senckenbeq,. biol. 49; pp 107-118. Nat. Hist.
BAERENDS, G.P. & J.M. BAERENDS-VAN ROON (1950) An intro- WICKLER, W. (1962a) Zur Stammesgeschichte funktionell
duction to the study of the ethology of cichlid fishes. Be- korrelierter Organ- und Verhaltensmerkmale: Ei-Attrappen
haviour suppl. I. 243 pp. und Maulbrüten bei afrikanischen Cichliden. Z. Tier-
BALON, E.K. (1977) Early ontogeny of Labeotropheus Ahl, psychol. 19; pp 129-164.
1927 (mbuna, Cichlidae, Lake Malawi) with a discussion WICKLER, W. (1962b) Egg-dummies as natural releasers in
on advanced protective systems in fish reproduction and mouth-breeding cichlids. Nature 194; pp 1092-1093.
development. Env. Biol. Fish. 2,”(2); pp 147-176. WICKLER, W. (1966) Über die biologischen Bedeutung des
BERNS, S., E.H. CHAVE & H.M. PETERS (1978) On the biology Genital-Anhangs der männlichen Tilapia macrochir.
of Tilapia squamipinnis (Günther) from Lake Malawi Senckenbeq, biol. 47; pp 419-422.
(Teleostei: Cichlidae). Amh. Hydnbiol. 84; pp 218-246.
FRYER, G. & T.D. ILES (1972) The Cichlid Fishes of the Great
Lakes of Africa. Oliver and Boyd, Edinburgh & London.
GOLDSCHMIDT, T. (1991) Egg mimics in haplochromine cich-
lids from Lake Victoria. Ethology. 88; pp 177-190.
GOLDSCHMIDT, T. & J. DE VISSER (1990) On the possible role of
egg-mimics in speciation. Acta Bioth. 38; pp 125-134.
HEINRICH, W. (1967) Untersuchungen zum sexualverhaltung
der Gattung Tilapia (Cichlidae, Teleostei) und bei
Artbastarden. Z. Tierpsychol. 24; pp 684-754.
HERT, E. (1989) The function of egg-spots in an African mouth-
brooding cichlid fish. Anim. Behav. 37; pp 726-732.
JACKSON, P.B.N. & A.J. RIBBINK (1975) Mbuna. TFH Publ.,
New Jersey. (Illustr, in colour) 128 pp.
KAPRALSKI, A. (1990a) Breeding Pseudotropheus tropheops.
TFH June. pp 86, 89-92. Pseudotropheus tropheops OB morph (Chinyamwezi Reef).
KAPRALSKI, A. (1990b) Spawning Labeotropheus fuelleborni. The TA-position: the male’s snout contacts the female’s body
TFH July. pp 10-13, 17 illustr. near the vent while she deposits some eggs.

96
 2. The spawning-techniques of Malawian haplochromines
Ad Konings

All the haplochromine species of Lake Malawi are It is therefore not enough to say “spawning by the usual
maternal mouthbrooders. Usually spawning begins haplochromine method”. Moreover a second spawn-
with the manoeuvre known as circling with alternat- ing variant (more advanced?), whereby the TA-posi-
ing T-positions (TA-TB: see Trewavas, this vol. p. 93) tion is not practised, has been described (see Konings,
which brings the pair to a synchronised spawning act 1989).
when the eggs are laid, fertilized, and taken into the This second technique is employed by (probably) a
mouth almost simultaneously, While the female de- large group of non-mbuna. I have observed it in C.
posits the egg(s) the male has its head close to the borteyi, Maravichromis lateristriga, Protomelas
female’s vent (TA-position). This position is very char- taeniolatus and Copad. sp. “Kawanga”. During the
acteristic for most mouthbrooders. All haplochromine entire spawning act the male never has its head near
mouthbrooders display the TB-position where the fe- the female’s vent (TA-position). In order to bring the
male collects the eggs and/or sperm. As Wickler pair to a synchronized spawning act the male leads
(1962a) recorded for his “H. wingatii”, some species the female to the spawning-site where an initial TB-
practise fertilization either before or after the female position culminates in a circling around each other.
has picked up the eggs. Recent descriptions by After a few rounds the male discontinues the circling.
Kapralski (1990) show that in some species behav- While the female discharges her eggs the male waits
iour is flexible in response to the environmental situa- beside her (passively?) until she has collected the eggs
tion. I have observed that females collect the eggs more in her mouth. Then he circles her again and quivers
rapidly in a crowded situation than in a peaceful one. his anal fin over the nest. Although quivering, the male
The fertilization of the eggs outside the female’s does not move forward but probably exudes its se-
mouth (external fertilization) has now been recorded men. The female snaps at the male’s vent and prob-
for several Malawian species, e.g. Dimidiochromis ably inhales the milt to fertilize the eggs inside her
compressiceps, Pseudotropheus sp. “Tropheops mouth. We may call this spawning technique the “to-
Chilumba” (Kapralski, 1990c, 1990a), Nimbochromis tal oral fertilization” method as all eggs are fertilized
livingstonii, Cyriocara moorii, and Labeotropheus inside the female’s mouth.
trewavasae (Konings, 1989). Most of these species Of the three fertilization techniques mentioned the
are known to fertilize (partly) the eggs in the mouth facultative oral fertilization method is the one most
as well, i.e. D. compressiceps (Knabe, pers. comm.), commonly observed in the aquarium. Facultative and
Labeotropheus spp. (Balon, 1977; Bailey, pers. total oral fertilization have both been observed in the
comm.), C. moorii (De Langhe, Pers. comm.), Ps. natural environment (pers. obs.: Ps. zebra and Ps. sp.
tropheops (pers. obs.). It seems that these species prac- “Elongatus Ornatus”, and Copad. sp. “Kawanga” re-
tise an oral fertilization when they feel disturbed. We spectively). For references see page 96.
may call this technique the “facultative oral fertiliza-
tion”. It is not known whether these conditions are Acknowledgements
natural or induced by the confines of the aquarium.
Many more species may practise an environmentally I gratefully acknowledge the discussions I enjoyed
induced variation of fertilization. with Dr. Ethelwynn Trewavas about many aspects of
So far, N. livingstonii is the only species of the group cichlid-ichthyology and with Gerard Tijsseling and
of which oral fertilization has not (yet) been observed. Peter Baasch about spawning behaviour in cichlids.

A pair in TB-position. Note the egg near the anal fin. The “total oral fertilization” method is practised by C. borleyi.

97
 CICHLID ORGANIZATIONS WORLDWIDE

Australia Taiwan (R.O.C.)

The New South Wales Cichlid Society Taiwanese Cichlid Association
P.O. Box 163 Nº17,Lane 239,An-Ho Road
Moorebank, N.S.W. 2170 Taipei

Queensland Cichlid Group United Kingdom

P.O. Box 163 British Cichlid Association
Wooloongabba, Queensland 4102 100 Keighley Road
Skipton, North Yorkshire, BD23 2RA
Victorian Cichlid Society
23 Mangana Drive U. S. A.
Mulgrave, Victoria 3170 American Cichlid Association
P.O. Box 32130
Austria Raleigh, NC 27622
Deutsche Cichliden Gesellschaft
Victor Kaplan Straße 1-9/1/3/12 Adv. Cichl. Aquarists South California
A-1220 Wien P.O. Box 8173
San Marino, CA 91108
Belgium Apistogramma Study Group
Belgische Cichliden Vereninging 1845 Jaynes Road
Kievitlaan 23 Mosinee, WI 64455
B-2228 Ranst
Cichlasoma Study Group
Czechoslovakia 6432 South Holland Court
SZCH Klub Chovatelov Cichlíd Littlerton, CO 80123
Príkopova 2
CS-831 03 Bratislava Fort Wayne Cichlid Association
9638 Manor Woods Rdf.
Denmark Ft. Wayne, IN 46804
Dansk Cichlide Selskab Greater Chicago Cichlid Association
Ornevej 58, st. tv. 2633 N. Rhodes River
DK-2400 Kobenhavn NV Grove, IL 60171

France Greater Cincinnati Cichlid Association

Association France Cichlid 15 W. Southern Avenue
15 Rue des Hirondelles Covington, KY 41015
F-67350 Dauendorf
Illinois Cichlids and Scavengers
Germany 7807 Sunset Drive
Deutsche Cichliden Gesellschaft Elmwood Park, IL 60635
Eberescheweg 41 Michigan Cichlid Association
D(W)-4200 Oberhausen P.O. Box 59
New Baltimore, MI 48047
Hungary
Hungarian Cichlid Association Ohio Cichlid Association
Lukács Laszló, Karolina ut 65 3896 Boston Rd.
H-1113 Budapest Brunswick, OH 44212

Netherlands Pacific Coast Cichlid Association

Nederlandse Cichliden Vereniging P.O. Box 28145
Boeier 31 San Jose, CA 95128
NL-1625 CJ Hoorn Rocky Mountain Cichlid Association
5065 W. Hinsdale Cir.
Sweden Littleton, CO 80123
Nordiska CiklidSällskapet
Skogsgläntan 16 Southern California Cichlid Association
S-435 38 Mölnlycke P.O. Box 574
Midway City, CA 92655
Switzerland
Deutsche Cichliden Gesellschaft Texas Cichlid Association
Am Balsberg 1 6845 Winchester
CH-8302 Kloten Dallas, TX 75231

98
4
 The CICHLIDS yearbook

Xenotilapia spiloptera, photographed by Ad Konings at Mabilibili, Tanzania.

Retroculus lapidifer from the Rio Tocantins. Photo by Ron Bernhard.

ISBN 3-928457-05-5 99