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CICHLIDS yearbook
Ad Konings (Ed.)
Volume 1
The
2
Cover photographs:
1 - Benthochromis tricoti, Lake Tanganyika, Zambia.
2 - “Cichlasoma” labridens, Media Luna, Mexico.
3 - Symphysodon aequifasciatus, variety.
4 - Otopharynx lithobates, Zimbawe Rock, Lake Malawi, Malawi.
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)
René Krüter (Krüter Tropicals, Rotterdam, Netherlands)
Roland Numrich (Mimbon Aquarium, Köln, Germany)
Edwin Reitz (Aquapport, Ronnenberg, Germany)
Dirk Verduijn (Verduyn Cichlids, Zevenhuizen, Netherlands)
1
2 3 4
Distributors:
USA: Old World Exotic Fish, Inc., P.O.Box 970583, Miami, Florida 33197
UK: Finz (U.K.), Ltd., Lady Ann Mills, Lady Ann Road, Batley, West Yorkshire WF17 0PS
Sweden: Fohrman Aquaristik AB, Pepparplan 4, 393 65 Kalmar
Germany: Aquapport (Edwin Reitz), Köselstraße 20, 3003 Ronnenberg
Netherlands: NVC, Lieshoutseweg 31, 5708 CW Stiphout
ISBN 3-928457-00-4
Copyright © 1991. Verlag Dr. Gertrud Dudin. 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.
Edition Cichlid Press
Verlag Dr. Gertrud Dudin, 6837 St. Leon-Rot, Germany
Printed by RAKET B.V., Pijnacker, Holland
3
Fascination with cichlids has taken many aquarists
to the point where they go and observe these won-
derful fishes in the natural habitat. The hunger for
more knowledge about the cichlids’ behavior and
distribution, and especially for more new species
has led to the introduction of an overwhelming ar-
ray of previously unknown species and varieties.
The speed with which new species are discovered
or re-discovered is sometimes too great to give a
newcomer a chance to establish itself in the hobby.
We, the authors of the yearbook, want to give a
chance to those species which are new or unknown
and which deserve a wide distribution among
hobbyists. Starting with this yearbook every year
the newest, the most important, the most interest-
ing and, not least, the most beautiful cichlids will
be introduced to the hobbyist. All (re-)introductions
will be accompanied by a color picture and the cur-
rently known information. Such information can
only be supplied by aquarists (or scientists) who
are specialists in their fields. Many authors have
contributed to the first edition of the cichlid year-
book. They will be introduced by the editor.
The yearbook starts with the section “Tangan-
yikan Cichlids”. René Krüter, who is importer of
Tanganyikan cichlids (Krüter Tropicals, Holland),
has made several expeditions to Lake Tanganyika.
He reports on his observations of Benthochromis
tricoti and some other species.
The second section, “Malawian Cichlids” is high-
lighted by an article written by Dr. Ethelwynn
Trewavas. She is the world’s most renowned cichlid-
ichthyologist and has worked for more than 60 years
on Malawi cichlids.
Photographer, diver and naturalist Mark Smith
(Fountain Valley, California) reflects his vivid in-
terest in cichlids in the various articles he has con-
tributed to the yearbook. Peter Baasch (Freiburg,
Germany) has bred several Malawi cichlids for the
first time in captivity. He reports on a beautiful
predator from the lake.
Victoria cichlids deserve a much better apprecia-
tion among aquarists than has been the case up to
now. Laif DeMason, who imports the widest vari-
ety of cichlids from all over the world (Old World
Exotic Fish, Miami, Florida), has made several ex-
peditions to Lake Victoria and knows these fish from
The CICHLIDS yearbook
first hand. His contribution deals with beautiful and
new discoveries.
The cichlids of West Africa are totally different
from the previous groups. Specialist Roland
Numrich (Mimbon Aquarium, Köln, Germany) has
made many expeditions to this part of the world and
has caught, bred and raised many of the newer finds.
His article describes two new species of Chromido-
tilapia.
The section “Central American Cichlids” starts
with an excellent article by Juan Miguel Artigas
Azas (San Luis Potosí, Mexico), who has more than
15 years experience of the natural habitats of Mexi-
can cichlids. On one of his countless expeditions
he re-discovered Paraneetroplus nebuliferum,
which had been elusive since its original descrip-
tion in 1860.
Willem Heijns (Stiphout, Netherlands) is editor
of the periodical of the Dutch Cichlid Association
and has kept and bred almost every Central Ameri-
can cichlid. He has contributed several fine reports
on the latest developments.
The section “South American Cichlids” has been
written by two esteemed specialists, Ron Bernard
(Rotterdam, Netherlands) and Frank Warzel (Mainz,
Germany). Ron has written many articles dealing
with South American cichlids and has collected
cichlids in Peru. Frank is super-specialized on
Crenicichla and knows more about the pike cichlids
than anybody else.
Gerard Tijsseling (Gouda, Netherlands) and John
Szwechlowicz (Holton-Le-Clay, UK) provide us
with valuable tips on how to better enjoy and breed
cichlids.
The yearbook ends with a literature section where
a few very important publications, related to
cichlids, are discussed. Nobody else could do this
better than the two authors Martin Geerts (Swalmen,
Netherlands) and Lee Finley (Pascoag, Rhode Is-
land). Martin is the ichthyological conscience of the
Dutch Cichlid Association and Lee that of the
American Cichlid Association.
This fantastic team of authors provides for an au-
thoritative treatment of the latest developments in
the cichlid-keeping hobby.
Your editor
Ad Konings (Editor)
4
In 1948 Poll described Benthochromis tricoti as
Haplotaxodon tricoti, but later, in 1984, he split the
two different populations, which he originally
thought belonged to one species, into two species:
Haplotaxodon tricoti and H. melanoides. Two years
later he gave these two species their own genus,
Benthochromis.
The name Benthochromis alludes to the assump-
tion that the species of this genus live at deep lev-
els. Benthos is Greek for “depth”. B. tricoti differs
from B. melanoides by its larger maximum size
TANGANYIKAN
CICHLIDS
The giant featherfin
René Krüter
A courting male Benthochromis tricoti, taken in Zambian waters by René Krüter.
which is about 20 cm; B. melanoides remains about
3 cm smaller. Also B. tricoti, at least in males, has
longitudinal stripes on the body. B. melanoides has
a number of black blotches on the head. The fe-
males of both species have an overall silvery-gray
coloration.
The food of B. tricoti consists of small inverte-
brates, such as crustaceans and zooplankton. Al-
though it attains a considerable size B. tricoti is not
a piscivore.
B. tricoti is mainly found in the southern part of
5
A male Benthochromis tricoti displaying.
When courting, Benthochromis tricoti has its mouth wide
open. This is also seen in species of Cyprichromis.
Lake Tanganyika. It is abundant in the deep to very
deep layers. Several investigations, including those
done by the Zambian Fisheries, revealed that the
densest populations were to be found between 100
and 150 meters deep. Another species which is
abundant at these levels is Cyphotilapia frontosa.
Normally I found B. tricoti where the rock for-
mations on shore descended at a steep angle. I have
never found this species in water shallower than 35
meters and at this level usually no more than a few
solitary individuals. At 65 meters I found many
more individuals, including several territorial males.
The population density between 100 and 150 me-
ters may be so high that many escape to higher lev-
els or they may migrate to higher levels to spawn.
Unfortunately it is impossible, with the available
diving equipment, to descend to deeper levels in
safety.
Sexually active males try to obtain a territory on
top of a conspicuous rock and defend this, like
Ophthalmotilapia ventralis, against conspecifics. I
have, however, never found sand in these territo-
ries. The bare rock is defended against rivals. Dur-
ing territorial disputes males show their finest col-
oration; the spectacle of fighting and courting B.
tricoti is one of the most fascinating splendors of
the lake.
The courting behavior is very characteristic. As
soon as a female approaches the breeding colony,
she is surrounded by a few males. Each of them
tries to lead her — while swaying his body heavily
— to his rock. When the male arrives at the rock he
takes a peculiar posture. With all fins erect the male
Benthochromis puts himself in an oblique, head-up
position. The dorsal fin is fully erected so that the
first few spines point forward. Also the mouth is
opened completely and the buccal sac is inflated.
The male remains in this position for a few seconds
showing his best to the female. Sometimes a weak
vibration enhances the display.
Unfortunately I have never witnessed a spawn-
ing. The size of the fry in the mouth of some fe-
males is remarkable. Sometimes the fry had a length
between 3 and 4 cm. This may indicate that the lar-
vae and fry feed inside the female’s mouth. If they
feed within the mouth the female has to eat appro-
6
A male Benthochromis tricoti leading a female (not visible) to its nest.
priate food, or she must release the fry from time to
time to let them eat by themselves. This may fur-
ther indicate that the female takes care of the fry
even after they have been released for the first time.
Successful breeding in captivity may give an an-
swer to these questions.
B. tricoti is an ideal aquarium resident. It is nei-
ther aggressive nor predatory. Despite its relatively
large dimensions, B. tricoti does not require an enor-
mous aquarium. After a relatively short acclimati-
zation period males regain their splendid colora-
tion. This in contrast to e.g. Cunningtonia and
Cyathopharynx, who need a long period of accli-
matization before full coloration returns.
B. tricoti is one of the most difficult to catch fish
in Lake Tanganyika. The main reason is the depth
of their preferred habitat. At just a few known loca-
tions they can be found at depths of less than 40
meters. Yet 40 meters is rather deep for a Scuba
diver. If we want to collect fish as well we encoun-
ter some problems. A method has to be developed
to bring these fish to the surface alive. It is fatal to
take a cichlid straight from these depths to the sur-
face. The closed swim-bladder, which is filled with
air, will expand drastically because of the decreased
pressure at the surface and may even burst. The
other internal organs will be forced out via the anus
and throat, which leads to the inevitable death of
the fish. That is why cichlids from these depths have
to be decompressed very carefully, in order to give
the blood enough time to absorb the excess air from
the bladder. The time it takes to surface B. tricoti,
caught at 35 meters, is five days.
I have noticed that fish can voluntarily alter their
depth range much quicker than when they are forced
to do it. My observation is that decompression takes
much longer when a fish is brought to the surface
against its will.
Another problem with B. tricoti is the females.
When we descend to a depth of 40 meters all
Benthochromis will be scared away. The territorial
males will return after a while, but of the females
no sign at all. The females seem to stay in small
schools far away from anything dangerous. The few
females that have been caught up to now got acci-
dentally stuck in the net.
Personally speaking, I think B. tricoti is one of
the most beautiful aquarium fishes. It has a very
interesting behavior and I hope to solve a few of its
mysteries on my future trips to the lake.
7
The cichlids from Lake Tanganyika are well-known
for their diverse behavior and enjoy a wide interest
among hobbyists. Of particular interest are the so-
called sand-cichlids. These fish, mainly of the ge-
nus Xenotilapia, live over the open sandfloor of the
lake. To protect them against predators their bodies
are silvery or sand colored and they live in large
schools, which gives protection to the individual.
During the breeding season the males of at least
three sand dwelling species acquire a beautiful and
conspicuous breeding dress. One of these species,
Enantiopus sp. “Kilesa” has not been described yet;
the other two species are Enantiopus melanogenys
and Xenotilapia ochrogenys. The latter two species
have been observed in the natural habitat in Burundi,
Tanzanian, and Zambian waters, but are reportedly
also present along the western coast of the lake
(Poll, 1956). E. sp. “Kilesa” was exported from
Zaire, probably from a location named Kalumbie.
E. melanogenys was described by Boulenger in
1898 as Ectodus melanogenys. A few years later, in
1906, the same author placed it in Enantiopus. Poll
placed this cichlid in Xenotilapia (1951) and back
in Enantiopus (1986). X. ochrogenys was described
as Enantiopus ochrogenys by Boulenger in 1914 and
placed – by Poll (1946) – in Xenotilapia.
A male Enantiopus sp. “Kilesa” courting a female.
Three beautiful sanddwelling cichlids
Greenwood (1978) demonstrated clearly that both
these species possess different anatomical features
which means that they are unlikely to belong to the
same genus.
E. melanogenys and E. sp. “Kilesa” are very slen-
der, elongated cichlids with a long snout. E. sp.
“Kilesa” differs from E. melanogenys by the shorter
lower jaw (see photos). All known species of the
genus Xenotilapia have slender bodies but rounded
heads. Their snout profiles descend steeply. The two
species of Enantiopus reach a maximum size of
about 16 cm; the maximum size of X. ochrogenys
is about 12 cm.
There are no distinct geographical races known
for E. melanogenys nor for E. sp. “Kilesa”. The
populations of X. ochrogenys along the coast of
Burundi and Tanzania do not vary – regarding male
coloration – but in Cameron Bay, Zambia, another
variety (or species) has been observed. The so-
called “Ndole Ochrogenys” differs from the wide-
spread variant of X. ochrogenys by prominent black
spots on the flanks and by the larger maximum size
– approximately 14 cm.
These sanddwelling cichlids forage in large
schools in rather shallow water. The maximum
depth recorded for E. melanogenys is 40 m but most
Ad Konings
8
A male Enantiopus melanogenys defending its territory.
The lower jaw is shorter in E. sp. “Kilesa” (above) than in
specimens are seen in depths of less than 10m. When
not breeding males have the same sandy, silvery
coloration as females. It gives them optimum cam-
ouflage on the sandy bottom. Most individuals of
one school remain together in the same school
throughout their lives. A school is probably formed
at the moment of the simultanuous release by the
mouthbrooding females. The youngsters grow and
breed together until they die after about three years.
All three species are mouthbrooders in which
only the female incubates the eggs and larvae.
Breeding has been observed throughout the year but
the highest activity takes place during the rainy sea-
son – from December until May. During the rest of
the year most of the schools move around and for-
age in different areas. X. ochrogenys normally oc-
curs in small schools of up to 30 members, but the
schools of the two Enantiopus may number in the
hundreds. Because there are so many fish in one
school together they do not have to seek shelter
when they want to breed, as many other sand-
dwelling cichlids do. Since most members of the
school have the same age they all attain their re-
productive phase at about the same time. It seems
that X. ochrogenys has attached itself to the breed-
9
The territory of E. melanogenys is flat (above) whereas
A male Xenotilapia ochrogenys from the Ndole Bay population.
ing colonies of the Enantiopus since they are usu-
ally found breeding together.
During the breeding season males defend their
territories and try to persuade females to spawn in
their nests. The actual breeding may take place in
bouts which would last for less than a week. Dur-
ing this period the fish do not eat but concentrate
on spawning. The season begins when the males
start staking out their territories in the sand. A male
E. melanogenys digs a flat, saucer-shaped territory
with a diameter of about 60 cm. In the center of the
territory, he digs a small pit with a diameter of about
15 cm. This will be the nest in which the spawning
will take place. The territory of X. ochrogenys is
rather peculiar; it consists of three to eight turrets
built by heaping sand. They are put in a circle around
the spawning-site which is a round, saucer-shaped
pit with a diameter of approximately 10 cm. The
territory of E. sp. “Kilesa” is a very interesting
“mixture” of the former two types. In an aquarium,
a male will occupy a large area in which he makes
several nests! These nests are shallow pits with a
diameter of about 15 cm. Around these nests the
male heaps sand turrets all over its territory, some-
times more than 20. A spawning usually takes place
10
in a single nest but next time they may spawn in
another.
The nests of these sanddwelling cichlids are me-
ticulously cleared of small pebbles which are larger
than the prevailing sandgrains. The reason why
males take the trouble to remove larger grains is
obvious: the female will not then mistake a tiny peb-
ble for an egg when she collects them during spawn-
ing. On an evenly structured sand-nest the eggs are
more conspicuous and can be collected faster.
When the arena is ready the male starts courting.
Here we notice some differences between Enan-
tiopus and Xenotilapia. A male X. ochrogenys at-
tracts a female to the nest with its fins fully ex-
tended, except the first part of the dorsal fin which
is just half-extended. A male Enantiopus attracts
females in a most peculiar fashion. Rivals are chased
with all fins erect, but females are seduced to the
nest with all fins down! The only thing extended is
the buccal cavity as in Xenotilapia. Furthermore the
male lies almost completely on its side when at-
tracting the other sex. When more than one male is
around the female is not chased when she ignores
the courtship of a male. As a female passes over the
different territories we see the males, one by one,
lying on their sides and trying to seduce her. The
males, however, stay in their territories.
When a female is ready to spawn she responds to
the display of the male. As soon as she enters the
nest, the male circles around her with fins erect –
in Enantiopus as well as Xenotilapia. Males are so
excited that they first chase all other fish away from
the nest. Meanwhile, the female remains motion-
less in the nest. After clearing the site of intruders,
the male enters the nest with fins erect and buccal
cavity extended. He gently pushes the female in the
hind part of her body and so encourages her to start
circling around. During circling the male vibrates
his extended buccal cavity. After two to three unin-
terrupted rounds, the female suddenly slows down
and lay some eggs. The male too stops circling and
waits impatiently for the female to move from the
site and leave the eggs to be fertilized. One to eight
two-millimeter long eggs are deposited at a time.
As soon as the female moves forward the male
shoots over the eggs and releases its milt. The eggs
are thus fertilized outside the female’s mouth. At
the end of the spawning cycle the female will not
release any more eggs, but the ritual goes on long
after that. Every time she moves forward the male
shoots like an arrow over the barren sand. Some-
times a male can be so excited that it shoots over
the female when she does not move fast enough.
The batch sizes vary between 30 and 80 eggs for
the two species of Enantiopus and between 10 and
40 for X. ochrogenys. A male may spawn with two
females on one day. In a short period all females
are gravid. After three weeks the fry are released
and are rather small. In one year they are mature
and might spawn in the next breeding season.
During spawning, in captivity, the pair is fre-
quently disturbed by other fish. The male spends
much time chasing away intruders. It is possible
that the female gets upset by these frequent inter-
ruptions and she may need a break for half an hour.
Once I observed that the female continued spawn-
ing with another male, after such a break. On an-
other occasion I noticed that a badly disturbed pair
of E. melanogenys quit spawning and that the fe-
male spat out the few eggs some hours later. The
next day she continued spawning with the same
male.
E. melanogenys, E. sp. “Kilesa” and X. ochro-
genys are great residents for the Tanganyika
aquarium. We should not mix them with large or
rough fish which will very likely damage the some-
what fragile females. If we want to enjoy the spec-
tacular view of a male in full color, we must give
him a rival. Two or more males enhance each oth-
er’s activity and bring out the best colors. If only
one male is kept, with several females, spawning
takes place but usually without the sparkling colors
of the male.
References
KONINGS, A. (1988) Tanganyika Cichlids. Verduijn
Cichlids, Rotterdam, Netherlands.
POLL, M. (1956) Résultats scientif. expl. hydrob.
belge au lac Tanganika (1946-1947). Poissons
Cichlidae, vol. III, fasc. 5B
POLL, M. (1986) Classification des cichlidae du lac
Tanganika. Tribus, genres et espèces. Mémoires
de la Classe des Sciences. Acad. Roy. Belg. Coll.
in-8°-2e
série, T. XLV Fasc. 2.
11
In spring 1988, Walter Dieckhoff discovered a new
Lamprologine along the Tanzanian coast, near Ikola.
He managed to collect five specimens alive but even-
tually only two survived the strenuous trip to Europe.
These two fish happened to be a male and female. In a
bare tank they didn’t look much different from a vari-
ant of Neolamprologus brichardi, but Walter assured
me that they behave differently. He noticed that the
Cygnus, as I named them later, live in pairs or solitary
and not as gregariously living pairs which is known
for N. brichardi. Walter further observed that the
Cygnus remains close to the rocks—a little like N.
furcifer—and that they have a dark blue-brown col-
oration with conspicuous blue eyes. I was fortunate to
get this pair. I first placed them in a large tank with
other fish in order to get them acclimatized to the type
of water and to each other. The male and female har-
monized well together and I decided to put them to-
gether in a small breeding tank (50 litre) which was
decorated with rocks and a flower pot. Even at their
relatively large size of about 9 cm the pair did well.
After about two months I noticed the first fry. There
were three in total. With Cygnus it is not the quantity
but the quality that counts. These three babies were
the most gorgeous lamprologus I had ever seen. The
At a size of about 2 cm, a juvenile Neolamprologus sp. “Cygnus” shows its best coloration.
A wildcaught male Neolamprologus sp. “Cygnus”.
Neolamprologus sp. “Cygnus”
yellow-orange fins and patches on the head were ab-
solutely unique. About two weeks later the pair
spawned again – only one youngster made it to the
free swimming stage. The pair kept on spawning about
every fortnight for more than a year. The broods, how-
ever, never contained more than 16 fry. In females the
yellow coloration on the head remains much longer than
in males but is lost – except for a faint tinge – at a size of
about 5.5 cm. Tankraised Cygnus have larger nests, even
at a smaller adult size. When kept in a dimly lit tank, a
pair with 20 mini-butterflies is an unforgettable sight.
Ad Konings
12
Altolamprologus calvus is a small predator cichlid
which lives in the southwestern part of the lake. Its
maximum size is about 15 cm, but most adult indi-
viduals measure between 7 and 10 cm for females
and between 10 and 13 cm for males. The female –
at the same age as the male – is considerably smaller
than the male.
Three different geographical variants are known
for A. calvus. The holotype is from the population
with the deepest black coloration which is found in
the southern part of the Zairean coast and in
Cameron Bay. The White Calvus occurs around
Cape Chaitika which is probably the most eastern
population. Here it shares the habitat with the Yel-
low race of Altolamprologus compressiceps.
In 1988 René Krüter discovered a completely yel-
low race of A. calvus at Chilange Rocks. This loca-
tion lies between those of the black and the white
populations. Since then the Yellow Calvus has be-
come a valuable addition to the Tanganyika
aquarium.
The best way to keep Altolamprologus in a tank
is to restrict oneself to one pair only. At a size of
about 5 cm the Calvus can be sexed by inspecting
the vents. To be sure you should check several speci-
mens to get the idea how a female and male should
A wildcaught male Altolamprologus calvus from Chilange Rocks, Zambia.
Altolamprologus calvus (Poll, 1978)
look. Females, which are much smaller than males,
have a distinctly broader genital papilla than males.
The larger the fish the easier it becomes to “vent”
them. The chosen pair are best put together with
other fish in a community aquarium. You can keep
either A. calvus or A. compressiceps in one tank but
avoid having both species together.
When A. calvus is not breeding, it does not show
any territoriality. Breeding starts when a female
starts guarding an empty shell. Females have been
observed breeding in a shell in the lake as well. It
is important to use a small shell. I have spawned
several pairs of A. calvus on a monthly basis using
an empty Lanistes nyassanus shell. This is the snail
from Lake Malawi. In their natural habitat, A. calvus
normally choses a small pocket in a rock, especially
when the female is to big to fit in a Neothauma shell.
The male should not be able to enter the shell.
During spawning, when the female sticks her eggs
to the inside of the shell, the male will be over the
entrance of the shell and discharges its milt. The best
sign that the pair have spawned is the observation that
the female stays, most of the time, inside the shell.
The male will be territorial now and defend the shell
with great vigor. About two weeks after spawning the
first fry appear in the opening of the shell.
Ad Konings
13
Neolamprologus sexfasciatus occurs in the south-
ern half of Lake Tanganyika. It is mainly found, in
shallow water, in the intermediate biotope where
sand is present between the small heaps of rocks.
N. sexfasciatus occurs solitary or in pairs. Breed-
ing pairs normally have youngsters which they de-
fend until they produce the next spawn. Sometimes
juveniles of a size larger than 3 cm are still guarded
by the parents.
Three different geographical races are known.
The holotype was collected on the Zairean coast
and probably looked like the fish in the picture
above. The first variety that was exported came from
Zambia (see photo). It occurs along the entire Zam-
bian coast except for the eastern part near the Tan-
zanian border. The populations along the Tanzanian
coast are completely yellow and have been exported
several times.
The Golden Sexfasciatus, as this race is called,
has been successfully bred in captivity but the Blue
Sexfasciatus from Zambia never.
In behavior (as far as one can judge from obser-
vations in the aquarium) there is a difference be-
tween these two races too. The Blue Sexfasciatus
shows a much more piscivorous appetite than the
Golden variant. The former cannot be kept with
A wildcaught female Neolamprologus sexfasciatus from an unknown location in Zaire.
Neolamprologus sexfasciatus from Zambia.
Neolamprologus sexfasciatus (Trewavas & Poll, 1952)
fishes smaller than 5 cm! The race from Zaire seems
to have a similar behavior to the Golden Sex-
fasciatus.
It is impossible to keep more than one pair in an
aquarium. In very large tanks (more than 750 liter)
two or three females might do well, but usually they
quarrel among each other.
N. sexfasciatus is a cave-brooder and we there-
fore have to supply the pair with some caves. The
best way to proceed is to let the female adapt to her
new environment first. When she is acclimatized
then the male can be introduced.
Ad Konings
14
Neolamprologus leloupi was described by Poll (in
1948) from a single specimen which was collected
a little north of Moba, on the Zairean coast. Thirty
years later, the same author described Neolampro-
logus caudopunctatus, a cichlid from the Zambian
coast. This fish closely matches N. leloupi in ana-
tomical features but lacks the distinct black margin
in the tail fin. In 1988 Walter Dieckhoff discovered
two other populations of Leloupi-like cichlids on
the Tanzanian coast. One of these cannot be distin-
guished from the holotype of N. leloupi. The other
one closely resembles the Zambian N. caudo-
punctatus.
Pierre Brichard (1989) concludes in his monu-
mental book that N. leloupi occurs in different geo-
graphical populations and that N. caudopunctatus
should be regarded as a synonym for N. leloupi.
Recently Janicki imported another variant from
the shores of Zaire. A female of this race is shown
in the picture above. The trade name, under which I
acquired these pretty fish, is “Red Fin Caudo-
punctatus”. I think that Brichard’s opinion will be
accepted by most taxonomists, so we should call
this little cichlid Neolamprologus leloupi.
N. leloupi lives in the intermediate habitat, where
rocks lie scattered over a sandy bottom. In Zam-
A female Neolamprologus leloupi showing a defensive posture.
Neolamprologus leloupi (Poll, 1948)
bian waters it is one of the most abundant cichlids
in this biotope. Schools of hundreds of individuals
are a common sight.
In the aquarium, N. leloupi behaves rather peace-
fully as long as we keep just one pair. Most of the
male’s aggression is directed towards conspecific
males. It can be kept with very small fish too. N.
leloupi is mature at an age of about one year and
may spawn at a size of about 4.5cm. Although it
has been observed spawning in an empty Neo-
thauma shell, the pair prefers something larger than
that. If no cave is present they will start digging
one under a stone. During the process the entrance
to the cave is concealed by a wall of sand (or gravel).
If we provide the pair with a small flowerpot, with
its bottom removed and placed upsidedown in the
sand, we can chose the position of their territory.
They will readily accept the pot and try to conceal
it by throwing sand against it. The result is a heap
of sand with only the top of the pot sticking out.
The size of the territory depends on the size of the
fish and the population density of the aquarium, but
usually it is about 30 cm in diameter. Any intruder
is chased from this area but youngsters (to a size of
at least 2.5 cm) from a previous spawn are toler-
ated.
Ad Konings
15
Neolamprologus mustax is not a common cichlid,
either in the wild or in captivity. Its first introduc-
tion, in the mid-seventies, was hampered by a wrong
identification and by the fact that a fake coloration
was applied on a picture of the Cameron Bay vari-
ant, the so-called “Blue Mustax”.
The first specimens exported to Europe were
caught in Cameron Bay. The fish of this population
have a grayish-yellow body color and yellow fins.
But this race displayed mainly the rather dull gray
color when held in dealer’s tanks. Nevertheless, it
was named “Blue Petricola”. It was then still an
undescribed species but some thought it could be
N. petricola.
N. mustax is reported only from the southern part
of the lake where it has been found in Zambian
waters. It is a small, rock dwelling cichlid with a
maximum size of approximately 9 cm. Several dif-
ferent populations are known. Most of them have
fish with yellow fins. Some populations, like those
at Cape Chaitika and near Mpulungu, have com-
pletely yellow or orange-yellow individuals. The
name Mustax means beard which reflects the white
coloration on the chin and lower part of the head
and which is seen in all known populations.
A wildcaught Neolamprologus mustax, collected near Mpulungu, Zambia.
Neolamprologus mustax (Poll, 1978)
In its habitat N. mustax is difficult to observe as
it mainly lives in the dark cracks between the rocks.
It is thus also difficult to collect which makes it a
rather expensive Tanganyikan cichlid.
N. mustax belongs to a group of rock dwelling
Lamprologines in which we also find the better
known N. leleupi, N. longior and N. cylindricus.
All of these species live solitary or in pairs when
adult. They are rather intolerant of the presence of
conspecifics. It is therefore difficult to accommo-
date several individuals in one aquarium. It is a good
idea to acquire just one male and a female. If the
tank is large enough (about 750 liters with lots of
rockwork) both may be introduced at once but it is
better to let the female acclimatize first.
If not properly fed or kept, N. mustax may lose
its deep yellow-orange coloration. It needs a lot of
carotene-rich food like Cyclops, Mysis or other al-
gae-eating crustaceans. Besides a lot of shelter in
the rockwork it needs clean water with a high pH,
preferably above 8.0. The colors of N. mustax may
fade initially but after treating it right, it will dis-
play the golden yellow as brightly as in the wild.
Ad Konings
16
Greenwoodochromis christyi was observed under
water for the first time in 1988. Walter Dieckhoff
found a few individuals in the Zambian part of the
lake. He collected a few specimens of which only a
female survived transportation back to Europe. One
year later some christyi were exported by Chris
Blighnaut in Mpulungu.
Greenwoodochromis christyi belongs to a group
of Tanganyikan cichlids which have a coloration
pattern consisting of a few horizontal rows of iri-
descent scales. A well-known cichlid from this
group is Limnochromis auritus. Previously, all these
cichlids were placed in the genus Limnochromis.
The species involved are L. auritus, L. abeelei, L.
staneri, Greenwoodochromis christyi, G. bellcrossi
and Gnathochromis permaxillaris (see next page).
The difference between the species in Green-
woodochromis and those that remain in Limno-
chromis is that the former have more than 48 scales
in an horizontal row between the gill-cover and the
tail. The species in Limnochromis have a maximum
40 scales in a row. In view of the recent revision of
the Malawi cichlids, where the genera are distin-
guished mainly by their basic coloration pattern, it
A female Greenwoodochromis christyi, collected in Zambian waters.
Greenwoodochromis christyi (Trewavas, 1953)
would not surprise me if all these species were
placed back in their original genus in future.
The two species currently in Greenwoodochromis
are distinguished from each other by the size of the
eye and the shape of the mouth. G. christyi has a
relatively smaller eye than G. bellcrossi. The latter
also has a steeply inclined mouth, while that of G.
christyi is only moderately inclined.
Greenwoodochromis christyi has been observed
in the intermediate habitat at somewhat deeper lev-
els. René Krüter found most individuals at a depth
of 25 meters and deeper. Sometimes a pair was ob-
served while guarding their offspring. Spawning has
not been observed but it is probably similar to that
of L. auritus. This means that G. christyi is a
mouthbrooder where both female and male take care
of their offspring.
Greenwoodochromis christyi has proven to be a
rather difficult cichlid to keep in an aquarium. A
single specimen can be kept, without problems, with
other species of similar size but a “pair” need a very
large tank. All their aggression is directed towards
conspecifics. The solution to this problem could be
to let the female adapt to the tank first.
Ad Konings
17
One of the most remarkable cichlids of Lake Tan-
ganyika is Gnathochromis permaxillaris. Its most
interesting features are the very large mouth which
opens in a peculiar way (see photo on the backcover)
and the shape of the upper lip. It is a rather large
cichlid which is known to grow to a size over 18’cm.
Gnathochromis permaxillaris lives over muddy
substrates and is usually found deeper than 40 me-
ters. Although the enormous mouth may suggest
otherwise, mostly very small organisms constitute
the staple food of G. permaxillaris. These tiny in-
vertebrates live close to the bottom, sometimes in
the upper layer of the mud. The wide gape of
permaxillaris spreads the inflow of water, going into
the mouth when the gill covers are extended, over a
relatively large area. This weakens the flow so that
only small and light particles are carried in. The
large cichlid has to “vacuum clean” the muddy bot-
tom continuously in order to obtain enough mate-
rial to live on.
Gnathochromis permaxillaris is probably the most interesting cichlid from Lake Tanganyika.
Gnathochromis permaxillaris (David, 1936)
The same feeding technique is observed in the
aquarium where G. permaxillaris feeds predomi-
nantly from the bottom. It takes a while before it
adapts to picking up food in mid-water.
Gnathochromis permaxillaris is now regularly
exported from Zambia albeit in very small numbers.
It seems to be a very fragile fish to collect but when
acclimatized to the aquarium it proves to be a hardy
and undemanding fish. A very welcome factor –
completely in contrast to Greenwoodochromis
christyi – is its remarkably peaceful attitude towards
conspecifics.
The sexes can be identified by looking at the ven-
tral apertures; the genital opening of the female is
noticeably larger than that of the male. Unfortu-
nately, there are usually not enough specimens to
compare and two males or two females might hide
in the same cave without showing any hostility to-
wards each other.
Ad Konings
18
The cichlids of the genus Trematocara belong to the
deepest living fishes of Lake Tanganyika. Representa-
tives of this genus were frequently captured at a depth
of 200 meters. It is assumed that these cichlids mi-
grate freely between the very deep layers and the sur-
face layers of the lake. Especially at night they move
to shallower water. This means that a special construc-
tion of the swimbladder allows the fish to compen-
sate for the enormous difference in pressure. During
the daytime Trematocara lives in the depths probably
together with its food. It is known that a small crusta-
cean, a kind of shrimp, undergoes a diurnal vertical
migration as well. Masses of this shrimp are found in
shallow water at night.
Evolution has permitted Trematocara nigrifrons to
follow these crustaceans during their daily migrations.
This means that these cichlids live in permanent dark-
ness. They have well developed adaptations to this
circumstance; T. nigrifrons possesses a very large eye
and an extremely sensitive lateral organ on the head
with which it can register minute movements in the
water. The sensory pore system, which is present on
the head of other cichlids as well, is greatly enlarged
on the head of Trematocara. The organ, which con-
sists of pressure sensitive tubules, is visible as small
“holes” below the eye. In the aquarium, I noticed that
A male Trematocara nigrifrons collected in Zambian waters.
Trematocara nigrifrons Boulenger, 1906
food is only accepted (greedily) when it moves. Fro-
zen food must be stirred by other fishes before T.
nigrifrons eats it. I have seen it hovering above the
food on the sand, in a manner similar to Aulonocara,
and waiting for a movement to occur.
Roger Bills, who is a researcher and collecter of
Zambian fishes, managed to catch and ship ten speci-
mens of T. nigrifrons. Unfortunately the fish proved
to be very sensitive and, after one year, only three sur-
vived. Surprisingly I succeeded in breeding them. The
male is considerably smaller than the female and, when
excited, shows black and white edges to the fins and a
black throat. Regrettably I did not observe the spawn-
ing; I first knew of it when I noticed the female with a
mouthful of pale yellow eggs. After three weeks the
fry were about 8 mm in size, when the female spat
them out because of a stress caused by netting her out
of the tank. The approximately 40 fry immediately
ate Artemia and now, three months later, they are about
4 cm long and am I the proud owner of a school of T.
nigrifrons.
T. nigrifrons is not a colorful addition to any
aquarium but for a true Tanganyika fancier, who usu-
ally looks further than colors alone, it is a very inter-
esting and elegant species.
René Krüter
19
The specimens depicted in the photograph above are
only tentatively identified as Microdontochromis
tenuidentatus. The holotype of this species shows, al-
beit in preserved condition, a horizontal, mid-lateral
row of black spots. In some preserved specimens these
spots appear as vertical bars. The cichlids in the pic-
ture above show only such pattern when they are ex-
cited. On the other hand, their morphology and their
typical mouth strongly suggest that these fish are
conspecific with M. tenuidentatus.
The holotype, together with about 50 other speci-
mens, was collected in Zaïre near Vua. The fish in the
picture were collected in Zambia in Cameron Bay,
which is not far from Vua. Sporadically, Fishes of
Burundi has exported some specimens which also
matched M. tenuidentatus in coloration even when they
were not excited. These were doubtless collected in
Burundi waters.
It is known that Cameron Bay harbors some varie-
ties (or even distinct species) which are not found at
any other place in the lake, for example Petrochromis
trewavasae, the so-called Tiger Nasutus or the Ndole
Bay variety of Xenotilapia ochrogenys. It is therefore
possible that the nondescript cichlid, when not excited,
is a geographical variant of M. tenuidentatus which is
Two wildcaught specimens of Microdontochromis tenuidentatus of unknown sex.
Microdontochromis tenuidentatus (Poll, 1951)
only to be found in Cameron Bay.
Microdontochromis tenuidentatus is a maternal
mouthbrooder. It has been bred in captivity by Volker
Puttberg, Dinslaken, Germany. He noticed that the eggs
were rather large and that a female broods only a few
eggs at the time, although about 20 are deposited dur-
ing a spawning. The male does not seem to make a
nest or defend a territory. The eggs are brooded for
about four weeks before the fry are released. During
this period the female continues feeding like she has
nothing in her mouth not to be swallowed.
In the lake M. tenuidentatus lives in large schools
in very shallow water. Most of them are found in the
vicinity of aquatic weeds. The mouth is quite
protrusible and the upper arm of the upper jaw bulges
the skin of the snout. The head is laterally compressed,
which is an important characteristic of this species. It
seems that M. tenuidentatus feeds from the plankton
in the open water as well as from invertebrates which
live among the weeds.
The teeth in the outer jaws are minute and look rather
fragile. The lower pharyngeal bone is very thin and
slenderly shaped. The teeth on this bone are minute
and pointed. The diet of M. tenuidentatus must con-
sists of very soft invertebrates and or tiny organisms.
Ad Konings
20
Cichlids from the genus Petrochromis are easily
recognized by the structure of the mouth. The nu-
merous teeth in the large jaws are permanently vis-
ible in most species. Petrochromis orthognathus is
the only known species in the genus in which the
teeth are less numerous and where the teeth are in-
visible when the mouth is closed. A cichlid with
the trade name “Petrochromis Tricolor” was ex-
ported from Zambia. The specimen which I was able
to have a closer look at showed just a few rows of
teeth in the mouth. It could close the mouth com-
pletely, “hiding” all teeth. The coloration pattern
closely matches that of P. orthognathus, i.e. a brown
upper half of the body and a lighter lower half.
Petrochromis polyodon occurs in several differ-
ently colored populations of which at least five have
been exported. The cichlid we have previously
called Petrochromis famula is the polyodon race
from Kigoma. Petrochromis polyodon is easily rec-
ognized by the short lower jaw and the generally
brown coloration of the females. Petrochromis
macrognathus also has a short lower jaw but its
mouth is underslung.
The “Red Texas Petrochromis” is closely related to
the “Texas Petrochromis” from the Tanzanian coast,
but is collected on the Zairean side of the lake.
A Zairean race of Petrochromis polyodon which is named “Kaiser Petrochromis” in the trade.
Above: The “Red Texas Petrochromis”; below:
Petrochromis, three new variants
Ad Konings
21
Many different populations of Tropheus moorii are
known and many more will be found in the future.
The genus Tropheus consists of at least six species
of which one is scientifically undescribed. This spe-
cies, named T. sp. “Black” or—previously—T. sp.
aff. brichardi, lives in the northern half of the lake,
whereas T. moorii is restricted to the southern half.
Although both species have been regarded as
conspecific by several authors, Walter Dieckhoff
discovered a locality where both species live
sympatrically (Kibwesa, Tanzania).
A wildcaught male Tropheus moorii, probably from Vua, Zaïre.
Tropheus moorii from an unknown location in Zaïre.
Tropheus moorii Boulenger, 1898
Ad Konings
In Zambia and along the lower half of the Zaïrean
coast the populations of T. moorii can be split into
two different groups. The holotype of T. moorii was
collected near Mpulungu and belongs to the group
of light-colored races which are further character-
ized by a colored region on the center of the body.
In Cameron Bay and north of it, populations of the
second group are found. This group consists of dark-
colored races which also have a colored cheek. There
is no location known where individuals of both groups
live together which would have indicated that we are
dealing with two different species.
Although there is a rather large difference, in col-
oration, between the two groups (see photographs),
they probably belong to the same species. The two
groups therefore represent two main geographical
variants. The Kasaba Bay in Zambia might have
separated these two groups for a long period of time
and they might thus have developed, independently
from each other, into the present races. The north-
ern borderline between the two groups is not known
but a dark-colored variant has allegedly been col-
lected near Vua in Zaïre. Further north into Zaïre
races of the light-colored group have been found.
One of them is shown in the smaller picture.
22
Cyathopharynx furcifer is a rather common and
spectacularly colored cichlid from Lake Tangan-
yika. Males attain a total length of about 18 cm
maximum; females, usually, are considerably
smaller. Cyathopharynx furcifer lives mainly over
sandy areas in shallow water. However, such sandy
bays must be close to rocky shores. The reason why
they prefer the rocks nearby is unknown because
they predominantly feed from the sandy substrate.
The food consists of diatoms which are collected
from the upper layer of the sediment on the sand or
on the rocks. Large quantities of material are
scooped up and processed through the long intesti-
nal tract. While feeding, the faeces can be seen
growing from the vent. A lot of the material swal-
lowed is indigestible. This results in an easy pas-
sage of the food through the guts. We must keep
this fact in mind when maintaining this cichlid in
the aquarium. Cyathopharynx furcifer needs a lot
of food but only a little meat.
In the lake, males congregate into large breeding
colonies. Each male makes its own nest which is a
kind of sand-castle with a diameter of approximately
50 cm. The nests are about 30 cm high. The differ-
ent territories can be just two meters apart. Breed-
This race of Cyathopharynx furcifer lives on the Ruziba reef in the northeastern part of the lake.
Cyathopharynx furcifer (Boulenger,1898)
ing colonies of C. furcifer are a spectacular sight;
sometimes more than 100 colorful males are found
together defending their nests. The females remain
in large schools, normally close to the rocky coast.
Because C. furcifer seems to be dependent on
areas with rocks, which are not present along the
entire coast uninterruptedly, it has developed sev-
eral geographical races. The Ruziba race, which is
shown in the photo, was exported from Burundi a
few years back.
When C. furcifer is collected the fishermen catch
the most beautiful males. These males keep their
colors for just a few weeks. Rarely do they regain
their full palette of colors in captivity, even though
they might spawn and produce offspring. Tankraised
males, however, show their best colors when still
small and usually keep them until something large
or rough disturbs them in the aquarium. In the
aquarium they need a large area with plain sand
where they can build their nest. Only in very large
tanks (over 750 liters) can we perhaps have more
than one male in color. The presence of a second,
smaller, male usually results in the most dominant
male retaining his colors.
Ad Konings
23
Xenotilapia flavipinnis is a common, sanddwelling
cichlid which is found all round the lake. It is found
predominantly over sand at depths between 1 and
30 meters. X. flavipinnis lives in schools. The larg-
est schools are found in the very shallow water.
Foraging schools may number over a hundred indi-
viduals. Like all other members of Xenotilapia it
feeds by sifting the sand. The food consists mainly
of soft-bodied invertebrates, like insect larvae.
It seems that X. flavipinnis has traditional forag-
ing and breeding grounds and that it is thus restricted
to certain areas. This has probably led to the exist-
ence of geographical races. Three of these races
have been exported to date. The holotype belongs
to the northern race of Magara, Burundi. The yel-
low variant from Nyanza Lac, Burundi, is shown in
the picture. From Zambia a variant is frequently
exported which shows a few rows of yellow scales
on the flank.
Xenotilapia flavipinnis breeds in colonies but it
is not a lek-breeder. Pairs separate from the school
– the school splits up into many breeding pairs –
and establish a rather large territory. Even though
we speak of a breeding colony the pairs are usually
more than two meters apart. For a small cichlid –
the maximum size of X. flavipinnis is about 10 cm
The yellow Xenotilapia flavipinnis from Nyanza Lac.
Xenotilapia flavipinnis Poll, 1985
– this is a large territory. In order to avoid territo-
rial disputes in the aquarium it is therefore best kept
as a pair.
Spawning occurs at any site in the territory. The
site might change during one spawning. There is
no visible nest constructed but it seems that the pair
chooses a slight dip in the sand. A few days before
the actual spawning, the male courts the female by
positioning himself with all fins erect and the buc-
cal cavity extended. The female answers in a simi-
lar way. Without much introductory behavior the
female starts laying the eggs which are fertilized
by the male while they are still on the sand. After
the female has taken all the eggs in her mouth she
stays with the male in the territory. After about 9
days the larvae are spat on the sand, in front of the
male who picks them up immediately. The male
carries the larvae for another 5 to 6 days before he
releases the fry in the territory. The fry are tended
by both parents – mostly by the male – for another
week or two. A further spawning usually occurs
within a month in the same territory.
Xenotilapia flavipinnis is a rather peaceful cichlid
which is best kept with non-predatory cichlids in
an aquarium with fine sand on the bottom.
Ad Konings
24
Xenotilapia papilio was described from a popula-
tion 40 kilometers south of Moba on the Zairean
coast. The two individuals shown on this page very
likely belong to the same species and are thus geo-
graphical variants. X. papilio is a very small
sanddwelling cichlid. Its maximum size lies around
8 cm. This very pretty species is mainly character-
ized by the underslung and tiny mouth. The teeth
are bicuspid, which excludes them from the genus
Asprotilapia which is characterized by tricuspid
teeth. However, X. papilio closely resembles A.
Several geographical variants of Xenotilapia papilio occur in the southeastern part of Lake Tanganyika. This race is
known as the “Sunflower Xenotilapia”.
Xenotilapia papilio. This variant is known as “Chituta Bay
Xenotilapia”.
Xenotilapia papilio Büscher, 1990
leptura and is also found sympatrically with the
latter.
Xenotilapia papilio lives in shallow water but at
some locations it may penetrate into depths of more
then 30 meters. In the deeper areas is is commonly
found in intermediate habitats, where sand and rocks
are both present. In very shallow areas it occurs over
pure sand. Large schools have never been observed.
Usually pairs or small groups are sighted. Like X.
flavipinnis it pairs off and defends a territory. Ob-
servations of aquarium populations show that once
a pair is formed they stay together for several con-
secutive spawns. Xenotilapia papilio is best kept
as a pair in a community tank as long as there are
no predatory species accompanying them. They re-
gard similar looking species, such as X. flavipinnis,
as competitors which elicit frequent quarrels.
Spawning in X. papilio is preceded by fin-dis-
plays by both male and female. One or two days
later the female deposits the eggs on a site some-
where in the territory. These are fertilized by the
male while they are still on the sand. The female
initially broods the eggs. The male takes the larvae
after about 8 to 10 days. After less than a week the
fry are released.
Ad Konings
25
Xenotilapia sp. “Katete” looks similar to the recently
described X. papilio but definitely is another species.
It has a shorter snout, a larger eye, a differently shaped
dorsal fin and a shallower caudal peduncle. The maxi-
mum length of X. sp. “Katete” lies around 7 cm, a
little bit smaller than X. papilio. In contrast to latter
species, the dorsal fin of X. sp. “Katete” is entirely
orange in color. The spines of the anterior part are
longer than those of the rest of the fin. The body has a
faint blue hue caused by a few rows of blue-reflecting
scales. Personally, I find this the most attractive spe-
cies among the small, rock-dwelling Xenotilapia.
The village of Katete lies near the border between
Zambia and Zaïre. In October 1988 we were looking
there for a good spot to collect Cyphotilapia frontosa.
While I was diving at 35 to 40 meters, I noticed X. sp.
“Katete” but at first glance I thought it was X.
flavipinnis. Realizing that that species is not commonly
found at these depths I looked closer and recognized
it from a picture from Horst Walter Dieckhoff in Ad
Konings’ book.
Xenotilapiasp.“Katete”inhabitstheintermediatezone
between sand and rocks, usually on a steeply sloping part
of the coast. C. frontosa and Neolamprologus buescheri
were among the other fishes of its habitat, which would
Xenotilapia sp. “Katete” lives at the deeper levels of the rocky habitat.
Xenotilapia sp. “Katete”
be a bad combination for the aquarium.
With a lot of effort we could only collect a small
number of these fragile cichlids of which just five sur-
vived the transport to my tank. They were three fe-
males and two males. There are no clear morphologi-
cal differences between the sexes. Unfortunately, X.
sp. “Katete” is very selective when it has to form pairs.
I was unlucky that none of the five specimens really
liked each other.As long as there had not been a spawn-
ing the pair seemed to be doing well; the other three
“Katetes” were then “expelled” from the 1000 liter
aquarium. These three were housed in a similar sized
tank of my friend Willem Bastinck.
Xenotilapia sp. “Katete” is a biparental mouth-
brooder, this means that the larvae are transferred to
the male after the female has brooded them for about
10 days. The problem with my pair started when the
male had taken the larvae in his mouth, because then
the female started to demolish the male, who reacted
by spitting out the larvae. Fortunately, Willem had
more success and was able to save some of the larvae
after the male had spat them out. Later, my pair also pro-
duced some fry which I was able to save from the fe-
male’s aggression. Now, after three years, the aquaristic
future of these beautiful cichlids looks a bit brighter.
René Krüter
26
Cyprichromis sp. “Leptosoma Jumbo”
Cyprichromis sp. “Leptosoma Jumbo”, a morph of the Kampemba race.
Cyprichromis sp. “Leptosoma Jumbo”, a yellow-tailed morph from an unknown location in Zaïre.
Ad Konings
27
The small cichlids from the open water in Lake Tan-
ganyika belong mainly to the genus Cyprichromis.
There are only two species described in this genus
but at least two others are known. The main differ-
ence between these species lies in the color patterns
of the males. They have gorgeous colors which are
clearly seen in their natural environment. Cypri-
chromis is spread over the entire lake but only C.
microlepidotus occurs in the northern part, north of
Nyanza Lac. From observations in Zambian waters
(Walter Dieckhoff; René Krüter, pers. comm.) it is
known that several species of Cyprichromis may
live in direct contact with each other.
Cyprichromis microlepidotus, which has a lake-
wide distribution, is characterized by small scales.
These are smaller than those from any other known
species of the genus. It is also larger than C.
leptosoma, which is a small and slenderly built
cichlid. The latter species differs from C. sp.
“Leptosoma Jumbo” by its shallow body and smaller
adult size only. C. sp. “Leptosoma Jumbo” can grow
to a size, at least under aquarium conditions, of ap-
proximately 12 cm whereas C. leptosoma rarely
exceeds a length of 9 cm. The fourth species, C. sp.
“Leptosoma Zebra”, is recognized by its yellowish
coloration – its lack of conspicuous colors – and by
the broad vertical bars which are not permanently
shown. C. leptosoma (Blue Flash, Isanga race), C.
microlepidotus, (Yellow Zambia race), C. sp.
“Leptosoma Jumbo” (Black Fin), C. sp. “Leptosoma
Zebra” and Paracyprichromis nigripinnis (Blue
Neon) have all been seen together in a small area in
Chitutu Bay.
C. sp. “Leptosoma Jumbo” is a pleasant aquarium
fish which normally shows its full colors. It is im-
possible to keep this species in pairs. Males actively
defend a territory and need the presence of at least
Cyprichromis sp. “Leptosoma Jumbo”, a morph from the
Kampemba population.
C. sp. “Leptosoma Jumbo”, a male of the yellow-tailed
morph from the Zaïrean race.
four other males to minimize the damage done to
the weaker one. Males behave rather peacefully to-
wards the females. In three breeding colonies, that
I have kept over several years, I have never found a
female killed by an aggressive male. Depending on
the number of males in the aquarium, they will stake
out their territories about 30 to 60 cm apart. The
males stay in mid-water and relate the size of their
premises to the distance from neighboring males.
Females are continuously courted and attracted to
the territory. When a female reacts to the male’s
invitation, the male bends its body and all its fins,
except the ventrals, away from the female. The ven-
tral fins, which have a large yellow spot, are held
to the outside, in front of the female’s mouth. In-
stinctively, the female snaps at the male’s ventral
fins. Since spawning takes place in mid-water, this
action might indicate to the female the spot where
she should deposit her eggs. The color of the spot
in the ventral fin does not exactly match that of the
egg. I have also observed males snapping at the pre-
sented fins of a more dominant male.
When a female is ready to spawn she remains
inside the male’s territory. Initially she will snap at
the male’s ventrals several times before she starts
depositing the eggs. He might discharge his milt
before the eggs are laid. At a certain moment the
male positions himself over the female, all fins ex-
tended, and pushes her gently, with his fully opened
mouth, on her head. The female releases one or a
few eggs and immediately backs up to retrieve it.
The sequence of release and backing up may be re-
peated several times before the female is led back,
by the male, to the center of the territory. Then fol-
low a number of “ventral-snappings” before the fe-
male discharges the next batch of eggs. The incu-
bation period lasts about three weeks.
28
Of all the new imports of the last years, the “Blue
Neon Cyprichromis” is the favorite of most Tan-
ganyika cichlidists. It is a small and peaceful cichlid
with a gorgeous coloration. The maximum size of
Paracyprichromis nigripinnis is about 10 cm for
aquarium specimens. Its shape closely resembles
that of P. brieni, the other species of the genus. The
difference lies mostly in their preferred habitat
depth, although P. nigripinnis has a noticably larger
eye (relatively seen). The latter is also found in
deeper regions of the rocky coast than P. brieni,
which lives in the upper 10 meters. The “Blue Neon
Cyprichromis” is caught at a depth between 20 and
30 meters. In Chituta Bay it lives in large caves or
in close vicinity of large boulders.
Paracyprichromis nigripinnis has a lake-wide
distribution but not all populations have such an
attractive pattern as the race shown in the photo-
graph. René Krüter (pers. comm.) found two dif-
ferent populations in Chituta Bay. The difference
between the two is the color of the dorsal fin, which
is more red in the other race. Walter Dieckhoff found
a race in Kigoma Bay which does not show the
“neon” stripes.
Paracyprichromis nigripinnis and P. brieni were
previously included in the genus Cyprichromis but
Paracyprichromis nigripinnis from Chituta Bay is known as “Blue Neon Cyprichromis”.
Paracyprichromis nigripinnis (Boulenger, 1901)
because of the difference in the number and arrange-
ment of the vertebrae they were placed in a new
genus. Although the splitting-up might seem a mat-
ter of opinion, P. nigripinnis and P. brieni are prob-
ably not at all related to the species currently in
Cyprichromis. In my opinion, they just show a re-
markable case of parallel evolution, whereby spe-
cies of two different ancestral origins have devel-
oped into similar looking species.
The most important distinction between these two
genera, however, lies in their spawning behavior.
The eggs of Cyprichromis are fertilized inside the
female’s mouth (see page 27) but those of Para-
cyprichromis are not.
Male P. nigripinnis defend a territory alongside
a large rock or inside a cave. Females are not at-
tracted to the nest by fin display but are seques-
tered from the school and then led to the male’s
territory. The female swims head-down, close to the
substrate, and discharges the eggs. As the eggs pass
by her head they are collected by the female. Mean-
while, the male stays above or beside the female
and discharges – visible clouds of sperm are occa-
sionally seen – its milt continuously. He fans the
milt towards the falling eggs which are thus ferti-
lized before the female picks them up.
Ad Konings
29
Of all the different biotopes Lake Malawi has to
offer, the rocky regions have brought forth the great-
est variety of colorful aquarium fishes. The much
less explored sandy areas may still hide many beau-
tiful species. Due to the openness of the sandy habi-
tats it is difficult, and sometimes impossible, to
collect some specimens. Since on the sand there is
no place to hide, almost all species living there have
a light, silvery-yellow coloration which camou-
flages them. Furthermore, the sanddwellers often
live in schools which gives them a protection against
Placidochromis phenochilus is a very conspicuous cichlid from the sand.
MALAWIAN
CICHLIDS
The blue sanddwellers
predators.
In contrast to most sanddwellers, there is a group
of conspicuously colored cichlids which usually live
solitarily or in very small groups. This group con-
sists of at least five different species. These have
no ancestor in common and do not belong to the
same genus. However, they have an important and
peculiar part of their feeding behavior in common
(see later).
The five conspicuously colored sanddwellers are
Cyrtocara moorii, Protomelas annectens, Otophar-
Ad Konings
30
Placidochromis electra with a digging Taeniolethrinops
preaorbitalis.
The distribution of Placidochromis electra is not restricted to Likoma Island. This specimen was pictured near Fort
ynx selenurus, Placidochromis electra, and
Placidochromis phenochilus. The first three species
have a lake-wide distribution. P. electra occurs
around Likoma Island and probably all along the
eastern shores, down to Fort Maguire. P. pheno-
chilus, which seems to be closely related to P.
electra, has been observed only in the northwest-
ern part of the lake. I found it near Mdoka, Chesese,
Chirwa Island, and between Selewa and Kasinda.
P. phenochilus is characterized by its white lips
and blue body. Even juveniles of 6 cm length have
the typical dark blue coloration. Its inclusion in
Placidochromis is based on its close resemblance
to P. electra. Of the about 25 individuals I have seen
at several locations, not a single one showed its
basic coloration pattern. The vertical barring, which
is faintly present in most individuals, is not a diag-
nostic feature since species from several other gen-
era show vertical bars – especially courting males
– as well.
The five species of the group have a particular
feeding strategy. They are attracted to stirred-up
material and usually follow a large sand-sifting spe-
cies like Taeniolethrinops preaorbitalis. The food
of most sanddwelling species consists of inverte-
brates which live and hide in the sand. There are
several ways to obtain these. Lethrinops-like spe-
cies dig in the sand and sift it through the gills.
Species of the genus Aulonocara detect their prey
by “listening to the sand”. Other species blow away
the upper layer in order to reveal the hiding inver-
tebrates. The five blue sanddwellers let other spe-
cies do the work. T. preaorbitalis is a large cichlid
and its continuous digging efforts create a lot of
stirred-up material. Although it seems that the blue
sanddwellers select their food from the clouds of
material spilled through the gills of the large sand-
sifter, they are actually more interested in search-
ing the ploughed sand for exposed invertebrates.
31
A male Protomelas annectens is rarely seen in its full blue breeding coloration. Photo taken near Otter Point.
Protomelas annectens is the most frequently seen follower of Taeniolethrinops preaorbitalis (photo taken near Fort
32
Otopharynx selenurus from Senga Bay
These crawl back into the sand as soon as possible.
The intriguing question is, why are these
sanddwelling species so conspicuously colored?
Especially P. phenochilus, O. selenurus and to a
somewhat lesser extent C. moorii, always show a
distinct blue coloration. These three species seem
to be mostly dependent on a large, sand-sifting host.
The other two species are also found foraging on
their own. It is therefore possible that their conspicu-
ous colors might warn off other followers – espe-
cially other species – from joining their host. In
sandy regions, cichlids commonly occur in groups
or schools, often of mixed species and it would thus
be normal for an individual to join such a group.
The amount of food revealed by the ploughing ac-
tivities of a large digger may not be sufficient for
more than one adult follower. If the follower is
highly dependent on the host, it may signal its po-
sition to the other species by taking on a territorial
coloration. In particular individuals of the same
species are wary of approaching an “occupied” host.
C. moorii and O. selenurus occur in very shal-
low water and are rarely found below depths of 10
meters. P. phenochilus and P. electra are usually
observed in water deeper than 15 meters whereas
P. annectens is found at any level.
The differences not only in the basic melanin
patterns but also in breeding behavior point to dif-
ferent ancestral origins for these sanddwelling
cichlids. The breeding behavior of C. moorii has
been observed several times by different aquarists.
They all noticed that as soon as the female had de-
posited some eggs they were fertilized by the male
while they were still on the sand. The same sequence
is also employed by P. annectens. Neither of the
two species build a nest or any structure to indicate
the spawning site. Although I have never observed
any of these blue sanddwellers spawning in their
natural environment, the courting behavior of the
males indicates that spawning can take place at any
site. Mouthbrooding and fry-guarding C. moorii and
P. annectens have been observed on the open sand.
P. electra is kept and bred by many hobbyists.
Spawning in the aquarium has revealed that the eggs
are fertilized inside the female’s mouth, as is the
case in most other Malawian cichlids. P. pheno-
chilus has yet not been exported as an aquarium fish.
Because P. phenochilus resembles P. electra to a
great extent it may have a similar breeding behavior,
i.e. that the eggs are fertilized inside the female’s
mouth.
O. selenurus is infrequently exported as Haplo-
chromis Nussae. All specimens are collected in
Senga Bay. It has been bred in captivity (Baasch,
pers. comm.) but details of the spawning sequence
are not known yet. The juveniles show the genus-
specific blotch pattern until they have reached a
length of approximately 7 cm. O. selenurus is a very
attractive and peaceful cichlid and makes a valu-
able addition to a Malawi community tank.
References
ECCLES, D.H. & TREWAVAS, E. (1989) Malawian
cichlid fishes. The classification of some
Haplochromine genera. Lake Fish Movies,
Herten, Germany.
DRUMMOND, B. (1976) How I keep.....Haplochromis
annectens. Bunt. Bull. (Am. Cichl. Ass.), Dec.,
pp. 29-30.
KOCHER, T.D. & MCKAYE, K.R. (1983) Defense of
heterospecific cichlids by Cyrtocara moorii in
Lake Malawi, Africa. Copeia (2), pp. 544-547.
KONINGS, A. (1989) Malawi cichlids in their natu-
ral habitat. Verduijn Cichlids, Rotterdam, Neth-
erlands.
SEIGARS, D. & BERARDO, T. (1979) Spawning the
Deep-Water Haplochromis. TFH, Vol. 27; April,
pp 4-12.
33
In May 1989, Walter Dieckhoff and I visited the
northern region of Lake Malawi, north of Chilumba
for the first time. Our trip took us to the Songwe
River which is the border between Malawi and Tan-
zania. It took us two days and several dives to con-
clude that there are no rocky reefs, coasts, or islets
between the border and Ngara. The entire coast,
here, consists of sandy and swampy beaches al-
though small rocks may lie scattered on the beach
and suggest that the area is rocky.
At Ngara, the most northerly rocky area in
Malawian waters, an intermediate type of habitat
supports a number of rock dwelling cichlids includ-
ing Aulonocara stuartgranti, the race with the or-
ange patch on the body. While diving there for the
first time I noticed some elongated sand-colored
cichlids which moved in small groups through the
habitat. At first glance they seemed to be of the
genus Aulonocara, a species of the sand dwelling
group. They were rather shy and I could take only
two photographs. At that time I did not see any in-
dividual with male coloration, which could be a fur-
ther indication that it belonged to the sand dwell-
ing group of Aulonocara. Of course I was eager to
have some specimens for a closer examination and
asked Saulos Mwale, our boat leader and an ex-
A male Aulonocara auditor patrolling its territory.
Aulonocara auditor (Trewavas, 1935)
tremely skilled fisherman, to catch a few. Saulos
needs only two words to describe the desired fish
and he will catch it for you, usually much better
colored specimens than the ones you had seen. Un-
fortunately, he could not find individuals with breed-
ing coloration which plainly meant they were not
there. However, two specimens were preserved and
they later turned out to be A. auditor.
At the next location where we dived, at Mdoka, I
found the same species again, but again no breed-
ing males. Further south, none of us saw A. auditor
again.
Gary Kratochvil and I went back to the same
places in December 1990, where I found A. auditor
in breeding coloration. In contrast to the few indi-
viduals I had seen in May 1989, we now observed
hundreds of them in large schools. Males in full
breeding coloration, however, were rare. Most in-
dividuals of this species of Aulonocara were found
below 15 meters of depth. The few territorial males
were found at about 20 m, at the deepest part of the
coast where rocks were still present.
The territorial behavior of the males resembles
closely that of the so-called Chitande Type Aulono-
cara; A. auditor might therefore belong to that group
of the genus as well. Males in breeding coloration
Ad Konings
34
Aulonocara ethelwynnae is found sympatrically with Aulonocara auditor.
Aulonocara auditor occurs in schools over sand near rocks. Aulonocara stuartgranti photographed at Mdoka.
“defend” their territories with little effort. Other
species and females are tolerated at all times within
the boundaries of the territory. Competing males are
“chased” by raising the dorsal fin and obstructing
their way. Under natural circumstances, I have never
observed fights, neither in A. auditor nor in Chitande
Type Aulonocara. In the aquarium, however, fights
may occur. Territorial males live rather far apart in
the wild and remain very static.
The territory of a male A. auditor usually lies on
the sand between some stones or rocks. They were
found only at the edge of the rocky habitat. Females
occur in large schools and forage over the sand.
Within such schools males with faint breeding
colors were observed. These males, sometimes
showed intolerance towards each other and “chased”
each other from the feeding site. No territories were
defended within the schools although the school and
its members remained rather static.
Other species of Aulonocara were found in the
schools together with A. auditor. Besides two dif-
ferent sand dwelling species, A. rostratum and A.
35
Aulonocara sp. “Chitande Type Kande”.
sp. “Jumbo Blue”, foraging individuals of A.
ethelwynnae (Northern or Chitande Aulonocara)
were also seen.
Ngara and Mdoka are the only two locations
where two different species of the Chitande Type
Aulonocara group are sympatric. Notwithstanding
the observation that A. auditor has a similar
behavior and habitat preference, it is morphologi-
cally clearly different from A. ethelwynnae. The
snout of A. auditor is considerably longer than that
of all other known species of the Chitande Type
Aulonocara. In all the other species the snout pro-
file is rounded whereas it is straight in A. auditor.
This might mean that A. auditor is able to poke its
snout deeper in the substrate than A. ethelwynnae
and thus feed from a different population of inver-
tebrates. Personally I think that food is abundant in
their environment and that those two species are
not competing with each other for food. The struc-
ture of the teeth in the jaws looks similar in all
Chitande Type Aulonocara; A. auditor is no excep-
tion. The four to six rows of teeth stand in a rela-
tively wide band anteriorly and become a single row
on the sides of the jaw. The flat lower jaw, with its
teeth, is somewhat reminiscent of that of several
Lethrinops species. A. auditor may be more of a
sand dwelling species than any other Chitande Type
Aulonocara. Territorial males of the other species
in this group are found throughout the year; they
don’t seem to have a particular breeding season. One
exception might be A. sp. “Chitande Type Kande”
breeding males of which were rare in June 1989. In
December 1990, this species was abundantly present
around Kande Island and many territorial males
could be observed.
At Kande Island, A. sp. “Chitande Type Kande”
shares the habitat with several other species of the
genus. Two of them, A. kandeensis and A. steveni,
are found in its immediate vicinity. It might thus be
possible that A. auditor and A. sp. “Chitande Type
Kande” breed mainly at a time when other species
of Aulonocara decrease their breeding activities.
These two species might be unsuccessful in obtain-
ing a territory when the other species are still at the
peak of their breeding season.
References
TREWAVAS, E. (1935) A synopsis of the cichlid
fishes of Lake Nyasa. Ann. & Mag. N. Hist. Ser. 10.
Vol XVI.
36
Abstract
The dark-bellied piscivorous cichlid of Lake Ma-
lawi is Tyrannochromis macrostoma (Regan), as it was
identified by Lewis et al. (1986: 39), who placed it
provisionally in the genus Cyrtocara. The darker lower
half of the head and body is usual for adults of this
species in the lake, but is masked by the blue colour
of breeding males and may be lost in unfavourable
circumstances. It is thus rarely seen in captive or pre-
served fishes.
The name T. nigriventer Eccles belongs to a differ-
ent species, related to Nimbochromis fuscotaeniatus.
The names T. maculiceps (Ahl) and T. polyodon
(Trewavas) may be synonyms of T. macrostoma but
this is not yet proved.
Introduction
Our friend Ad Konings, to whom we owe so much
for his books on the home life of the cichlids of
Lakes Tanganyika and Malawi, has contributed to
the solution of the problem of the specific identity
of the predator known as “Dark-belly” or “Schwarz-
bauch”. But in his article (Cichlidae (BCA) Vol. 11,
no. 3) he has made some conflicting statements that
A beautiful male Tyrannochromis macrostoma in breeding coloration. Photo by Stuart Grant.
The genus Tyrannochromis Eccles & Trewavas, 1989
Dr. Ethelwynn Trewavas
need clarification.
Konings writes (Cichlidae 11 (3): p. 73) “The so-
called Black-belly is thus correctly named T.
macrostoma”. But two paragraphs later “the species
currently known as T. macrostoma should be correctly
named T. nigriventer”.
These are apparently mutually contradictory state-
ments. In two publications that may be considered
“current”, namely Lewis et al. (1986; p. 39) and Ec-
cles & Trewavas (1989, p. 97. fig. 43), the name
macrostoma is correctly used for the species to which
Regan gave that name in 1922. Konings now tells me
that by “currently” he meant “currently in the
aquaristic trade”.Also in Konings’books, 1989, 1990,
macrostoma has been incorrectly used for nigriventer.
This is important for us to know because all the eco-
logical information we have about T. nigriventer is
found in Konings’books under the name macrostoma.
While the true macrostoma is there treated under the
name T. maculiceps, which may be a synonym of T.
macrostoma, or at least the name of a closely related
species or subspecies.
To explain how this confusion came about let me
relate the following, which also shows the value of
cooperation between field naturalists, aquarists and
taxonomists.
37
A male Tyrannochromis nigriventer in breeding coloration.
History of the identification of “Dark-belly”
The type species of the genus Tyrannochromis is T.
macrostoma, first described by Regan (1922) in the
genus Haplochromis on the basis of one specimen of
22.5 cm standard length (26 cm total length). Regan’s
figure is reproduced as fig. 43 in our book (Eccles &
Trewavas, 1989). The Christy collection contained five
other specimens of 12-20’cm total length. All these
have white flanks below the mid-lateral black stripe
and the markings above this stripe stand out on a pale
background. Dr. Digby Lewis and his collaborators
(1986; p. 39) have a striking photograph of a fish la-
belled Cyrtocara macrostoma with all the lower part
of the flank black or dark brown, and their text re-
marks on this peculiarity.
Could this be the same species as our white-bellied
museum specimens? We had verbal reports of the abil-
ity of this predator to change colour, but with no de-
tails about the conditions in which this might occur.
Then, when our book was nearly finished, Eccles
found in the Museum a specimen that had been
wrongly catalogued as H. polyodon. Its lower flanks
were tinged brown. Here, he thought, is our “Dark-
belly”, and as it was not either of the named species
of Tyrannochromis he described it as a “new species”
with the name T. nigriventer, which means “black-
belly”. The photograph of this fish is fig. 47 of our
book (Eccles & Trewavas, 1989). The chief measur-
able difference between this fish and the other species
of Tyrannochromis is the short upper arm of the upper
jaw (premaxillary pedicel). In T. nigriventer this is
less than one third of the length of the head. In the
other species it is very long, more than one third of
the length of the head, enabling the fish to protrude its
mouth as a long funnel without dislocating its jaws.
The photograph on p. 6 of Konings (1990a) shows the
mouth protruded, but not to its full extent. (This fish
is a male in which the overall blue colour masks the
adult pattern, of which the only indication consists of
a black mark on each scale, more prominent as the
ventral surface is approached, giving the flanks a soiled
appearance).
The paragraphs on p. 103 of Eccles & Trewavas on
coloration and ecology apply to T. macrostoma and
not to T. nigriventer.
So were Dr. Digby Lewis and his collaborators mis-
taken in identifying “Dark-belly” with T. macrostoma?
This question was answered by our diving naturalists.
First,Andreas Spreinat of Göttingen caught and pre-
served some “Dark-bellies” and also some other preda-
tors. These he sent to the BMNH for our opinions.
38
A juvenile Tyrannochromis nigriventer photographed at
Chewere. The tips of the premaxillary pedicels bulge the
skin clearly before the eyes. (Size approx. 10 cm TL)
Our book was then in the last stages, but after it was
published Gordon Howes made some measurements
for me. The pre-maxillary test showed that those la-
belled “Dark-belly” by Spreinat agreed with T.
macrostoma. The others were a good match for T.
nigriventer. Spreinat also found that in his photographs
the ends of the premaxillary pedicels could be seen
bulging the skin. His measurements of these, though
naturally inexact, gave a similar result, “Dark-belly”
having pedicels whose length went 2.0 to 2.6 times in
the head length, whereas those of the other species,
identified as N. fuscotaeniatus, went 3.5 to 3.75 times.
The relationship between N. fuscotaeniatus and T.
nigriventer will be mentioned below.
In November 1989 Ad Konings made a further visit
to Lake Malawi and he too caught a number of “Dark-
belly” and some of a species agreeing with T.
nigriventer. The premaxillary test again pointed to T.
macrostoma as the correct name for “Dark-belly”.
Both divers confirmed that the dark belly is the usual
live colour of adult, confident T. macrostoma, in the
lake (except blue breeding males). Konings (1989; p.
129) has a photograph of a guarding female that is
very dark except on the dorsal side, where the dorsal
and dorso-lateral series of markings stand out clearly
on a pale background. This photograph is captioned
T. maculiceps but Konings now believes, probably cor-
rectly, that it is T. macrostoma. The adoption of black
colouring when guarding young is reminiscent of the
colour changes in the mbuna Melanochromis auratus
and may well be a parallel phenomenon. Konings
states (1990a; p. 179) that the dark colour is not al-
ways present in individuals transferred to a tank.
Spreinat saw pale-bellied individuals that were “in-
disposed” (letter of 24-6-90). It is understandable that
a fish being caught for the pot or for preservation in a
museum is not merely “indisposed” but terrified.
Hence our pale-bellied museum specimens.
The problem of the specific identify of “Dark-belly”
is now therefore solved.
Konings touched on two other problems in the ge-
nus Tyrannochromis. The first is the validity of the
two other nominal species, T. polyodon (Trewavas,
1935) and T. maculiceps (Ahl, 1927). Each of these
was based on a single specimen. To synonymize T.
polyodon it is necessary to show that the greater depth
of its body comes within the range of variation of T.
macrostoma. To synonymize T. maculiceps it must be
shown that its narrow head and interorbital width, as
well as possibly its wider band of teeth at any given
size, all come within the range of a reasonable sam-
ple of T. macrostoma from the southern part of the
lake. The type specimen of T. maculiceps was caught
at the extreme northeastern part of the lake, a small
fish of 163 mm standard length. Other specimens as-
signed to this species are from Chilumba and Nkhata
Bay. It may be that this is a species or subspecies com-
mon in the north. Until these tests have been applied
the validity of the names remains an open question.
The second problem is the relationship between T.
nigriventer and Nimbochromis fuscotaeniatus. They
resemble each other in proportions and dentition, but
N. fuscotaeniatus has a pigment-pattern which is usu-
ally much less regular than in the figured holotype
(Regan, 1922 and Eccles & Trewavas, 1989; fig. 79),
forming irregular blotches as in other species of
Nimbochromis. No such tendency to form blotches is
seen in the holotype of T. nigriventer, in which the
ventero-lateral brown patch is broader and continu-
ous. Konings (1989; p. 222) remarks on the close re-
semblance between these two species.
Konings does not like to include fuscotaeniatus in
A juvenile Tyrannochromis macrostoma photographed at
Chitande Island. This specimen is identified by the fact
that the tips of the premaxillary pedicels bulge the skin
between the eyes. (Size approx. 8 cm TL.)
39
A female Tyrannochromis nigriventer defending her offspring. The photo was taken at Chinyankwazi Island. It shows that
a guarding female T. nigriventer can adopt a rather dark coloration on the belly. This female was collected for positive
identification.
Nimbochromis because of an alleged difference in
breeding habits (Konings, 1989; p. 128). The eggs of
N. livingstonii are fertilized on the substrate, then taken
into the mouth (Konings, 1989; p. 219), whereas those
of fuscotaeniatus are said to be fertilized in the mater-
nal mouth. The reference for the latter statement is a
paper by Dorenstouter (1982), but the identity of the
fish in the photograph accompanying this paper is
doubtful and the statement requires confirmation.
There are other differences between N. fuscotaeniatus
and typical species of Nimbochromis and the inclu-
sion of fuscotaeniatus requires further consideration.
References
DORENSTOUTER, C.F., (1982) Haplochromis fusco-
taeniatus Regan, 1921. NVC Periodiek (Dutch
Cichlid Ass.) April: 43.
ECCLES, D.H. & E. TREWAVAS, (1989) Malawian cichlid
fishes. The classification of some haplochromine
genera. Lake Fish Movies. Herten, West Germany.
335 pp.
KONINGS, A., (1989) Malawi cichlids in their natural
habitat. Verduijn Cichlids, Netherlands. 303 pp.
Illustr. in colour.
KONINGS, A., (1990a) Cichlids and all the other fishes
of Lake Malawi. TFH Publications, Inc. Neptune
City, New Jersey. 495 pp. Many illustr. in colour.
KONINGS, A. (1990b) The Malawi genus Tyran-
nochromis. Cichlidae (Brit. Cichl. Ass.) Vol 11 (3),
pp. 71-73.
LEWIS, D., P. REINTHAL, P. & J. TRENDALL (1986) Aguide
to the fishes of Lake Malawi national park. World
Wildlife Fund. Gland, Switzerland. 71 pp. illustr.
in colour.
SPREINAT,A. (1990) Zur Revision der Haplochrominen
des Malawisees nach Eccles & Trewavas. DATZ
(Aquar. Terrar. Z.) (4); pp 245-248, illustr.
This article originally appeared in the January 1991 issue
of Cichlidae, the periodical of the British Cichlid
Association. Details of the BCA may be obtained from
page 98.
40
Maravichromis (C.) liemi was previously placed in
Caprichromis Eccles & Trewavas (1989). The reason
was that it had a strongly inclined mouth, a broad lower
jaw and a specialized feeding behavior. Although it is
functional to split large genera into a few smaller ones,
monotypic or oligotypic genera should be avoided
when possible. The peculiar structure of the mouth of
liemi and orthognathus is related to the feeding
behavior but other species in the lake also have simi-
lar mouth structures and comparable behavior. In the
genus Protomelas there are several, undescribed spe-
cies (e.g P. sp. “Paedophage”) with a similar mouth
structure, but none of these or other species like
Diplotaxodon greenwoodi, have been put or will be
put into a new genus or into Caprichromis. Personally
I think the different mouth structure is insufficient justi-
fication for erecting a new genus for these two species.
M. (C.) liemi is reported to steal eggs and larvae from
mouthbrooding females (McKaye & Kocher, Animal
Behav. 31, 1983), specifically from P. pleurotaenia. The
authors probably meant Nyassachromis microcephalus
as P. pleurotaenia remains close to the bottom and
mouthbrooding females are not found in schools.
I have observed M. (C.) liemi at many different lo-
cations all round the lake and it seems likely that their
diet is not restricted to larvae and eggs alone. I have
A male Maravichromis (Caprichromis) liemi in full breeding coloration photographed at Eccles Reef.
A subadult Maravichromis (Caprichromis) liemi.
Maravichromis (Caprichromis) liemi (McKaye & McKenzie, 1982)
observed several times that M. (C.) liemi attacked male
cichlids as well, preferably those with parasites
(Argulus) on the throat region. It is therefore possible
that M. (C.) liemi feeds also, or even primarily, on these
parasitic invertebrates since fish with Argulus on their
throat are very common.
M. (C.) liemi is normally found near rocks whereas
M. (C.) orthognathus prefers the open sand. Breeding
males congregate into colonies and construct a sand-
castle nest (more than 1 m in diameter) against a rock.
Mouthbrooding females stay in schools and probably
release their fry in the rocky habitat.
Ad Konings
41
Sciaenochromis gracilis was described from three
specimens in the Christy collection. The total length
of the largest specimen measured 217 mm. Some sin-
gle specimens have been imported alive under trade
names such as “Torpedo” or “Dark Line Torpedo”.
Recently, the export of S. gracilis has taken a more
frequent character. This is certainly caused by the ef-
forts of Stuart Grant to comply with the increased de-
mand.
The distribution of this rare species seems to be re-
stricted to the southern part of the lake. Observations
about their feeding and breeding behavior and about
their breeding coloration are not published yet.
The specific name (gracilis = slender) is aptly cho-
sen. The narrowly pointed head with a slightly down-
ward curved profile is a striking feature that reminds
one of Nimbochromis linni.Aband, consisting of small
spots, runs diagonally from the nape to the upper part
of the tail (peduncle). In this respect it could be mis-
taken for S. spilostichus or Maravichromis formosus.
Maybe gracilis should be placed into Maravichromis
as has been suggested before (Konings, 1989).
Its large mouth, its teeth and its shape presume a
predatory lifestyle. The strong tail indicates a fast
swimmer with ability to spurt.
A courting male Sciaenochromis gracilis photographed in the author’s aquarium.
Sciaenochromis gracilis (Trewavas, 1935)
Outside the breeding period S. gracilis is a lively
swimmer. Every now and then they suddenly stop and
move their eyes noticeably. Although S. gracilis be-
haves rather peacefully we should not house them in
too small quarters. Their behavior changes during the
breeding period. Then S. gracilis is able to spurt over
large distances without apparent effort.
Courting is rather impetuous and intensive, but the
female is not hurt. Males defend a territory and con-
struct (in my tank) a crater nest with a diameter of
about 50 cm. The eggs are very small and are ferti-
lized inside the female’s mouth. Since the eggs are
small many fry can be expected. Females with a size
of approximately 15 cm may release over 100 fry. Ju-
veniles are, in comparison to other predators of simi-
lar size, very small and slim. They grow rather slowly,
probably because they show the same behavior as the
adults, i.e. they frequently stop moving and search the
area with their eyes only. Food must then be brought
into their mouths to let them eat! At a size of about 2-
3 cm they defend small feeding territories and quarrel
among themselves.
S. gracilis grows to a maximum size of about 25’cm
and is in shape, coloration, and behavior a valuable
addition to any large aquarium.
Peter Baasch
42
Otopharynx decorus is a small to medium sized cichlid
which is found predominantly over the open sandy bot-
toms of the lake. Its distribution encompasses probably
the entire lake because specimens have been collected at
Vua as well as in the southern part of the lake. The maxi-
mum size of O. decorus is about 18 cm (total length);
females are only a little bit smaller than males.
This sanddwelling species lives in small groups,
usually numbering not more than 6 members. Solitary
individuals are frequently observed, too. In Decem-
ber, when I made the observations, there was some-
times one male individual in the group that showed
some of its breeding coloration. Territorial males have
not been seen. Members of the genus Otopharynx usu-
ally spawn near rocks. It is therefore possible that O.
decorus also spawns close to a rocky coast and that
the males which I was able to observe were foraging
and not breeding.
The feeding behavior of O. decorus is rather char-
acteristic of an insectivorous cichlid. With its large eyes
it carefully screens the sand. Only after it has located
something interesting does it bite into the substrate.
Its food consists mainly of invertebrates which hide in
the sand. The teeth on the lower pharyngeal bone are
somewhat enlarged in the center, which is a common
A male Otopharynx decorus taken at a depth of approximately 18 meters in Senga Bay.
Otopharynx decorus (Trewavas, 1935)
feature in cichlids which live on crustaceans and in-
sects. The feeding behavior closely resembles that of
Maravichromis sp. “Double Spot” (see next page).
Both species have a small mouth with which they se-
lectively pick their food from the substrate. Both spe-
cies are found sympatrically.
The neutral coloration pattern consists of three rather
large blotches on the flank. These blotches are not al-
ways distinct. When observed in the somewhat sub-
dued light at a depth of 20 meters, the spots are almost
invisible. I have never seen them black as in O.
(Ctenopharynx) nitidus with which it is commonly
found sympatrically. The coloration of the body of O.
decorus is like that of the sand – light beige.
Among the sanddwelling cichlids of the lake it is
rather easily recognized by the combination of the
small mouth and the size of the fish. When observed
in its natural environment, its behavior will immedi-
ately allow an identification. Identification of aquarium
or preserved specimens is simplified by the fact that
O. decorus has a high number of soft rays in the dorsal
fin, 13 or 14, whereas most other Haplochromines have
less than 12. It has, furthermore, a high number of
scales on the flanks, higher than any other known Oto-
pharynx.
Ad Konings
43
Maravichromis sp. “Double Spot” is a rather small,
sand-dwelling cichlid. Its maximum size is known to
be less than 14 cm (total length). It has a rather wide
distribution since it has been found on the east coast
near Fort Maguire, as well as near Mdoka in the north-
ern part of the lake, and near Chintheche. It has, how-
ever, never been seen in Senga Bay, although cichlids
are frequently collected there.
M. sp. “Double Spot” is characterized by a diago-
nal row of three double-spots. In some specimens the
spots are so large that they form a broad, almost solid
black band. This feature helped me decide to put it in
the genus Maravichromis and not in Otopharynx.
The “Double Spot” prefers the open sandy regions
and is usually found at between 5 and 30 meters depth.
Most individuals forage on their own, rarely two or
three individuals are seen together. The coast near Fort
Maguire is inhabited by a rather dense population, al-
though they never forage in groups or schools. At the
two other locations where I have seen M. sp. “Double
Spot”, only single individuals were found. The fish of
the east coast population have a yellow coloration on
the body and especially on the fins, whereas speci-
mens from the other locations are more silvery and
have colorless fins.
Maravichromis sp. “Double Spot” scrutinizes the sand for something edible (Fort Maguire).
Maravichromis sp. “Double Spot”
The mouth of M. sp. “Double Spot” is very small
and is utilized to pick small invertebrates from the sand.
The substrate is therefore scrutinized for anything ed-
ible. The characteristic posture of this cichlid is poised
over the sand, visually screening the area in front of
the head with its large, movable eyes. The fish swims
from one place to the next and everywhere it halts and
scrutinizes the sand. In this respect M. sp. “Double
Spot” resembles Otopharynx decorus. The latter spe-
cies also forages in small groups.
The specialized feeding behavior of M. sp. “Dou-
ble Spot” might also be used for screening parasites
and fungus on the body and fins of sick fish. Once I
observed a female Nyassachromis prostoma, with a
visible fungus in the dorsal fin, who presented the ail-
ing fin by laying itself on the sand in front of a forag-
ing “Double Spot”. At the same moment a M.
labidodon disturbed the scene, so I could not see
whether M. sp. “Double Spot” would actually clean
the prostoma’s fin or not. It is, however, likely that it
does, otherwise the ailing fish would not present its
fin in the way it did. The specific coloration pattern of
the “Double Spot” is unique among the haplo-
chromines of the lake and might function as a signal
to other species that it is a “Cleaner”.
Ad Konings
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The Cichlids Yearbook Vol 1.pdf

  • 1. 1 CICHLIDS yearbook Ad Konings (Ed.) Volume 1 The
  • 2. 2 Cover photographs: 1 - Benthochromis tricoti, Lake Tanganyika, Zambia. 2 - “Cichlasoma” labridens, Media Luna, Mexico. 3 - Symphysodon aequifasciatus, variety. 4 - Otopharynx lithobates, Zimbawe Rock, Lake Malawi, Malawi. 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) René Krüter (Krüter Tropicals, Rotterdam, Netherlands) Roland Numrich (Mimbon Aquarium, Köln, Germany) Edwin Reitz (Aquapport, Ronnenberg, Germany) Dirk Verduijn (Verduyn Cichlids, Zevenhuizen, Netherlands) 1 2 3 4 Distributors: USA: Old World Exotic Fish, Inc., P.O.Box 970583, Miami, Florida 33197 UK: Finz (U.K.), Ltd., Lady Ann Mills, Lady Ann Road, Batley, West Yorkshire WF17 0PS Sweden: Fohrman Aquaristik AB, Pepparplan 4, 393 65 Kalmar Germany: Aquapport (Edwin Reitz), Köselstraße 20, 3003 Ronnenberg Netherlands: NVC, Lieshoutseweg 31, 5708 CW Stiphout ISBN 3-928457-00-4 Copyright © 1991. Verlag Dr. Gertrud Dudin. 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. Edition Cichlid Press Verlag Dr. Gertrud Dudin, 6837 St. Leon-Rot, Germany Printed by RAKET B.V., Pijnacker, Holland
  • 3. 3 Fascination with cichlids has taken many aquarists to the point where they go and observe these won- derful fishes in the natural habitat. The hunger for more knowledge about the cichlids’ behavior and distribution, and especially for more new species has led to the introduction of an overwhelming ar- ray of previously unknown species and varieties. The speed with which new species are discovered or re-discovered is sometimes too great to give a newcomer a chance to establish itself in the hobby. We, the authors of the yearbook, want to give a chance to those species which are new or unknown and which deserve a wide distribution among hobbyists. Starting with this yearbook every year the newest, the most important, the most interest- ing and, not least, the most beautiful cichlids will be introduced to the hobbyist. All (re-)introductions will be accompanied by a color picture and the cur- rently known information. Such information can only be supplied by aquarists (or scientists) who are specialists in their fields. Many authors have contributed to the first edition of the cichlid year- book. They will be introduced by the editor. The yearbook starts with the section “Tangan- yikan Cichlids”. René Krüter, who is importer of Tanganyikan cichlids (Krüter Tropicals, Holland), has made several expeditions to Lake Tanganyika. He reports on his observations of Benthochromis tricoti and some other species. The second section, “Malawian Cichlids” is high- lighted by an article written by Dr. Ethelwynn Trewavas. She is the world’s most renowned cichlid- ichthyologist and has worked for more than 60 years on Malawi cichlids. Photographer, diver and naturalist Mark Smith (Fountain Valley, California) reflects his vivid in- terest in cichlids in the various articles he has con- tributed to the yearbook. Peter Baasch (Freiburg, Germany) has bred several Malawi cichlids for the first time in captivity. He reports on a beautiful predator from the lake. Victoria cichlids deserve a much better apprecia- tion among aquarists than has been the case up to now. Laif DeMason, who imports the widest vari- ety of cichlids from all over the world (Old World Exotic Fish, Miami, Florida), has made several ex- peditions to Lake Victoria and knows these fish from The CICHLIDS yearbook first hand. His contribution deals with beautiful and new discoveries. The cichlids of West Africa are totally different from the previous groups. Specialist Roland Numrich (Mimbon Aquarium, Köln, Germany) has made many expeditions to this part of the world and has caught, bred and raised many of the newer finds. His article describes two new species of Chromido- tilapia. The section “Central American Cichlids” starts with an excellent article by Juan Miguel Artigas Azas (San Luis Potosí, Mexico), who has more than 15 years experience of the natural habitats of Mexi- can cichlids. On one of his countless expeditions he re-discovered Paraneetroplus nebuliferum, which had been elusive since its original descrip- tion in 1860. Willem Heijns (Stiphout, Netherlands) is editor of the periodical of the Dutch Cichlid Association and has kept and bred almost every Central Ameri- can cichlid. He has contributed several fine reports on the latest developments. The section “South American Cichlids” has been written by two esteemed specialists, Ron Bernard (Rotterdam, Netherlands) and Frank Warzel (Mainz, Germany). Ron has written many articles dealing with South American cichlids and has collected cichlids in Peru. Frank is super-specialized on Crenicichla and knows more about the pike cichlids than anybody else. Gerard Tijsseling (Gouda, Netherlands) and John Szwechlowicz (Holton-Le-Clay, UK) provide us with valuable tips on how to better enjoy and breed cichlids. The yearbook ends with a literature section where a few very important publications, related to cichlids, are discussed. Nobody else could do this better than the two authors Martin Geerts (Swalmen, Netherlands) and Lee Finley (Pascoag, Rhode Is- land). Martin is the ichthyological conscience of the Dutch Cichlid Association and Lee that of the American Cichlid Association. This fantastic team of authors provides for an au- thoritative treatment of the latest developments in the cichlid-keeping hobby. Your editor Ad Konings (Editor)
  • 4. 4 In 1948 Poll described Benthochromis tricoti as Haplotaxodon tricoti, but later, in 1984, he split the two different populations, which he originally thought belonged to one species, into two species: Haplotaxodon tricoti and H. melanoides. Two years later he gave these two species their own genus, Benthochromis. The name Benthochromis alludes to the assump- tion that the species of this genus live at deep lev- els. Benthos is Greek for “depth”. B. tricoti differs from B. melanoides by its larger maximum size TANGANYIKAN CICHLIDS The giant featherfin René Krüter A courting male Benthochromis tricoti, taken in Zambian waters by René Krüter. which is about 20 cm; B. melanoides remains about 3 cm smaller. Also B. tricoti, at least in males, has longitudinal stripes on the body. B. melanoides has a number of black blotches on the head. The fe- males of both species have an overall silvery-gray coloration. The food of B. tricoti consists of small inverte- brates, such as crustaceans and zooplankton. Al- though it attains a considerable size B. tricoti is not a piscivore. B. tricoti is mainly found in the southern part of
  • 5. 5 A male Benthochromis tricoti displaying. When courting, Benthochromis tricoti has its mouth wide open. This is also seen in species of Cyprichromis. Lake Tanganyika. It is abundant in the deep to very deep layers. Several investigations, including those done by the Zambian Fisheries, revealed that the densest populations were to be found between 100 and 150 meters deep. Another species which is abundant at these levels is Cyphotilapia frontosa. Normally I found B. tricoti where the rock for- mations on shore descended at a steep angle. I have never found this species in water shallower than 35 meters and at this level usually no more than a few solitary individuals. At 65 meters I found many more individuals, including several territorial males. The population density between 100 and 150 me- ters may be so high that many escape to higher lev- els or they may migrate to higher levels to spawn. Unfortunately it is impossible, with the available diving equipment, to descend to deeper levels in safety. Sexually active males try to obtain a territory on top of a conspicuous rock and defend this, like Ophthalmotilapia ventralis, against conspecifics. I have, however, never found sand in these territo- ries. The bare rock is defended against rivals. Dur- ing territorial disputes males show their finest col- oration; the spectacle of fighting and courting B. tricoti is one of the most fascinating splendors of the lake. The courting behavior is very characteristic. As soon as a female approaches the breeding colony, she is surrounded by a few males. Each of them tries to lead her — while swaying his body heavily — to his rock. When the male arrives at the rock he takes a peculiar posture. With all fins erect the male Benthochromis puts himself in an oblique, head-up position. The dorsal fin is fully erected so that the first few spines point forward. Also the mouth is opened completely and the buccal sac is inflated. The male remains in this position for a few seconds showing his best to the female. Sometimes a weak vibration enhances the display. Unfortunately I have never witnessed a spawn- ing. The size of the fry in the mouth of some fe- males is remarkable. Sometimes the fry had a length between 3 and 4 cm. This may indicate that the lar- vae and fry feed inside the female’s mouth. If they feed within the mouth the female has to eat appro-
  • 6. 6 A male Benthochromis tricoti leading a female (not visible) to its nest. priate food, or she must release the fry from time to time to let them eat by themselves. This may fur- ther indicate that the female takes care of the fry even after they have been released for the first time. Successful breeding in captivity may give an an- swer to these questions. B. tricoti is an ideal aquarium resident. It is nei- ther aggressive nor predatory. Despite its relatively large dimensions, B. tricoti does not require an enor- mous aquarium. After a relatively short acclimati- zation period males regain their splendid colora- tion. This in contrast to e.g. Cunningtonia and Cyathopharynx, who need a long period of accli- matization before full coloration returns. B. tricoti is one of the most difficult to catch fish in Lake Tanganyika. The main reason is the depth of their preferred habitat. At just a few known loca- tions they can be found at depths of less than 40 meters. Yet 40 meters is rather deep for a Scuba diver. If we want to collect fish as well we encoun- ter some problems. A method has to be developed to bring these fish to the surface alive. It is fatal to take a cichlid straight from these depths to the sur- face. The closed swim-bladder, which is filled with air, will expand drastically because of the decreased pressure at the surface and may even burst. The other internal organs will be forced out via the anus and throat, which leads to the inevitable death of the fish. That is why cichlids from these depths have to be decompressed very carefully, in order to give the blood enough time to absorb the excess air from the bladder. The time it takes to surface B. tricoti, caught at 35 meters, is five days. I have noticed that fish can voluntarily alter their depth range much quicker than when they are forced to do it. My observation is that decompression takes much longer when a fish is brought to the surface against its will. Another problem with B. tricoti is the females. When we descend to a depth of 40 meters all Benthochromis will be scared away. The territorial males will return after a while, but of the females no sign at all. The females seem to stay in small schools far away from anything dangerous. The few females that have been caught up to now got acci- dentally stuck in the net. Personally speaking, I think B. tricoti is one of the most beautiful aquarium fishes. It has a very interesting behavior and I hope to solve a few of its mysteries on my future trips to the lake.
  • 7. 7 The cichlids from Lake Tanganyika are well-known for their diverse behavior and enjoy a wide interest among hobbyists. Of particular interest are the so- called sand-cichlids. These fish, mainly of the ge- nus Xenotilapia, live over the open sandfloor of the lake. To protect them against predators their bodies are silvery or sand colored and they live in large schools, which gives protection to the individual. During the breeding season the males of at least three sand dwelling species acquire a beautiful and conspicuous breeding dress. One of these species, Enantiopus sp. “Kilesa” has not been described yet; the other two species are Enantiopus melanogenys and Xenotilapia ochrogenys. The latter two species have been observed in the natural habitat in Burundi, Tanzanian, and Zambian waters, but are reportedly also present along the western coast of the lake (Poll, 1956). E. sp. “Kilesa” was exported from Zaire, probably from a location named Kalumbie. E. melanogenys was described by Boulenger in 1898 as Ectodus melanogenys. A few years later, in 1906, the same author placed it in Enantiopus. Poll placed this cichlid in Xenotilapia (1951) and back in Enantiopus (1986). X. ochrogenys was described as Enantiopus ochrogenys by Boulenger in 1914 and placed – by Poll (1946) – in Xenotilapia. A male Enantiopus sp. “Kilesa” courting a female. Three beautiful sanddwelling cichlids Greenwood (1978) demonstrated clearly that both these species possess different anatomical features which means that they are unlikely to belong to the same genus. E. melanogenys and E. sp. “Kilesa” are very slen- der, elongated cichlids with a long snout. E. sp. “Kilesa” differs from E. melanogenys by the shorter lower jaw (see photos). All known species of the genus Xenotilapia have slender bodies but rounded heads. Their snout profiles descend steeply. The two species of Enantiopus reach a maximum size of about 16 cm; the maximum size of X. ochrogenys is about 12 cm. There are no distinct geographical races known for E. melanogenys nor for E. sp. “Kilesa”. The populations of X. ochrogenys along the coast of Burundi and Tanzania do not vary – regarding male coloration – but in Cameron Bay, Zambia, another variety (or species) has been observed. The so- called “Ndole Ochrogenys” differs from the wide- spread variant of X. ochrogenys by prominent black spots on the flanks and by the larger maximum size – approximately 14 cm. These sanddwelling cichlids forage in large schools in rather shallow water. The maximum depth recorded for E. melanogenys is 40 m but most Ad Konings
  • 8. 8 A male Enantiopus melanogenys defending its territory. The lower jaw is shorter in E. sp. “Kilesa” (above) than in specimens are seen in depths of less than 10m. When not breeding males have the same sandy, silvery coloration as females. It gives them optimum cam- ouflage on the sandy bottom. Most individuals of one school remain together in the same school throughout their lives. A school is probably formed at the moment of the simultanuous release by the mouthbrooding females. The youngsters grow and breed together until they die after about three years. All three species are mouthbrooders in which only the female incubates the eggs and larvae. Breeding has been observed throughout the year but the highest activity takes place during the rainy sea- son – from December until May. During the rest of the year most of the schools move around and for- age in different areas. X. ochrogenys normally oc- curs in small schools of up to 30 members, but the schools of the two Enantiopus may number in the hundreds. Because there are so many fish in one school together they do not have to seek shelter when they want to breed, as many other sand- dwelling cichlids do. Since most members of the school have the same age they all attain their re- productive phase at about the same time. It seems that X. ochrogenys has attached itself to the breed-
  • 9. 9 The territory of E. melanogenys is flat (above) whereas A male Xenotilapia ochrogenys from the Ndole Bay population. ing colonies of the Enantiopus since they are usu- ally found breeding together. During the breeding season males defend their territories and try to persuade females to spawn in their nests. The actual breeding may take place in bouts which would last for less than a week. Dur- ing this period the fish do not eat but concentrate on spawning. The season begins when the males start staking out their territories in the sand. A male E. melanogenys digs a flat, saucer-shaped territory with a diameter of about 60 cm. In the center of the territory, he digs a small pit with a diameter of about 15 cm. This will be the nest in which the spawning will take place. The territory of X. ochrogenys is rather peculiar; it consists of three to eight turrets built by heaping sand. They are put in a circle around the spawning-site which is a round, saucer-shaped pit with a diameter of approximately 10 cm. The territory of E. sp. “Kilesa” is a very interesting “mixture” of the former two types. In an aquarium, a male will occupy a large area in which he makes several nests! These nests are shallow pits with a diameter of about 15 cm. Around these nests the male heaps sand turrets all over its territory, some- times more than 20. A spawning usually takes place
  • 10. 10 in a single nest but next time they may spawn in another. The nests of these sanddwelling cichlids are me- ticulously cleared of small pebbles which are larger than the prevailing sandgrains. The reason why males take the trouble to remove larger grains is obvious: the female will not then mistake a tiny peb- ble for an egg when she collects them during spawn- ing. On an evenly structured sand-nest the eggs are more conspicuous and can be collected faster. When the arena is ready the male starts courting. Here we notice some differences between Enan- tiopus and Xenotilapia. A male X. ochrogenys at- tracts a female to the nest with its fins fully ex- tended, except the first part of the dorsal fin which is just half-extended. A male Enantiopus attracts females in a most peculiar fashion. Rivals are chased with all fins erect, but females are seduced to the nest with all fins down! The only thing extended is the buccal cavity as in Xenotilapia. Furthermore the male lies almost completely on its side when at- tracting the other sex. When more than one male is around the female is not chased when she ignores the courtship of a male. As a female passes over the different territories we see the males, one by one, lying on their sides and trying to seduce her. The males, however, stay in their territories. When a female is ready to spawn she responds to the display of the male. As soon as she enters the nest, the male circles around her with fins erect – in Enantiopus as well as Xenotilapia. Males are so excited that they first chase all other fish away from the nest. Meanwhile, the female remains motion- less in the nest. After clearing the site of intruders, the male enters the nest with fins erect and buccal cavity extended. He gently pushes the female in the hind part of her body and so encourages her to start circling around. During circling the male vibrates his extended buccal cavity. After two to three unin- terrupted rounds, the female suddenly slows down and lay some eggs. The male too stops circling and waits impatiently for the female to move from the site and leave the eggs to be fertilized. One to eight two-millimeter long eggs are deposited at a time. As soon as the female moves forward the male shoots over the eggs and releases its milt. The eggs are thus fertilized outside the female’s mouth. At the end of the spawning cycle the female will not release any more eggs, but the ritual goes on long after that. Every time she moves forward the male shoots like an arrow over the barren sand. Some- times a male can be so excited that it shoots over the female when she does not move fast enough. The batch sizes vary between 30 and 80 eggs for the two species of Enantiopus and between 10 and 40 for X. ochrogenys. A male may spawn with two females on one day. In a short period all females are gravid. After three weeks the fry are released and are rather small. In one year they are mature and might spawn in the next breeding season. During spawning, in captivity, the pair is fre- quently disturbed by other fish. The male spends much time chasing away intruders. It is possible that the female gets upset by these frequent inter- ruptions and she may need a break for half an hour. Once I observed that the female continued spawn- ing with another male, after such a break. On an- other occasion I noticed that a badly disturbed pair of E. melanogenys quit spawning and that the fe- male spat out the few eggs some hours later. The next day she continued spawning with the same male. E. melanogenys, E. sp. “Kilesa” and X. ochro- genys are great residents for the Tanganyika aquarium. We should not mix them with large or rough fish which will very likely damage the some- what fragile females. If we want to enjoy the spec- tacular view of a male in full color, we must give him a rival. Two or more males enhance each oth- er’s activity and bring out the best colors. If only one male is kept, with several females, spawning takes place but usually without the sparkling colors of the male. References KONINGS, A. (1988) Tanganyika Cichlids. Verduijn Cichlids, Rotterdam, Netherlands. POLL, M. (1956) Résultats scientif. expl. hydrob. belge au lac Tanganika (1946-1947). Poissons Cichlidae, vol. III, fasc. 5B POLL, M. (1986) Classification des cichlidae du lac Tanganika. Tribus, genres et espèces. Mémoires de la Classe des Sciences. Acad. Roy. Belg. Coll. in-8°-2e série, T. XLV Fasc. 2.
  • 11. 11 In spring 1988, Walter Dieckhoff discovered a new Lamprologine along the Tanzanian coast, near Ikola. He managed to collect five specimens alive but even- tually only two survived the strenuous trip to Europe. These two fish happened to be a male and female. In a bare tank they didn’t look much different from a vari- ant of Neolamprologus brichardi, but Walter assured me that they behave differently. He noticed that the Cygnus, as I named them later, live in pairs or solitary and not as gregariously living pairs which is known for N. brichardi. Walter further observed that the Cygnus remains close to the rocks—a little like N. furcifer—and that they have a dark blue-brown col- oration with conspicuous blue eyes. I was fortunate to get this pair. I first placed them in a large tank with other fish in order to get them acclimatized to the type of water and to each other. The male and female har- monized well together and I decided to put them to- gether in a small breeding tank (50 litre) which was decorated with rocks and a flower pot. Even at their relatively large size of about 9 cm the pair did well. After about two months I noticed the first fry. There were three in total. With Cygnus it is not the quantity but the quality that counts. These three babies were the most gorgeous lamprologus I had ever seen. The At a size of about 2 cm, a juvenile Neolamprologus sp. “Cygnus” shows its best coloration. A wildcaught male Neolamprologus sp. “Cygnus”. Neolamprologus sp. “Cygnus” yellow-orange fins and patches on the head were ab- solutely unique. About two weeks later the pair spawned again – only one youngster made it to the free swimming stage. The pair kept on spawning about every fortnight for more than a year. The broods, how- ever, never contained more than 16 fry. In females the yellow coloration on the head remains much longer than in males but is lost – except for a faint tinge – at a size of about 5.5 cm. Tankraised Cygnus have larger nests, even at a smaller adult size. When kept in a dimly lit tank, a pair with 20 mini-butterflies is an unforgettable sight. Ad Konings
  • 12. 12 Altolamprologus calvus is a small predator cichlid which lives in the southwestern part of the lake. Its maximum size is about 15 cm, but most adult indi- viduals measure between 7 and 10 cm for females and between 10 and 13 cm for males. The female – at the same age as the male – is considerably smaller than the male. Three different geographical variants are known for A. calvus. The holotype is from the population with the deepest black coloration which is found in the southern part of the Zairean coast and in Cameron Bay. The White Calvus occurs around Cape Chaitika which is probably the most eastern population. Here it shares the habitat with the Yel- low race of Altolamprologus compressiceps. In 1988 René Krüter discovered a completely yel- low race of A. calvus at Chilange Rocks. This loca- tion lies between those of the black and the white populations. Since then the Yellow Calvus has be- come a valuable addition to the Tanganyika aquarium. The best way to keep Altolamprologus in a tank is to restrict oneself to one pair only. At a size of about 5 cm the Calvus can be sexed by inspecting the vents. To be sure you should check several speci- mens to get the idea how a female and male should A wildcaught male Altolamprologus calvus from Chilange Rocks, Zambia. Altolamprologus calvus (Poll, 1978) look. Females, which are much smaller than males, have a distinctly broader genital papilla than males. The larger the fish the easier it becomes to “vent” them. The chosen pair are best put together with other fish in a community aquarium. You can keep either A. calvus or A. compressiceps in one tank but avoid having both species together. When A. calvus is not breeding, it does not show any territoriality. Breeding starts when a female starts guarding an empty shell. Females have been observed breeding in a shell in the lake as well. It is important to use a small shell. I have spawned several pairs of A. calvus on a monthly basis using an empty Lanistes nyassanus shell. This is the snail from Lake Malawi. In their natural habitat, A. calvus normally choses a small pocket in a rock, especially when the female is to big to fit in a Neothauma shell. The male should not be able to enter the shell. During spawning, when the female sticks her eggs to the inside of the shell, the male will be over the entrance of the shell and discharges its milt. The best sign that the pair have spawned is the observation that the female stays, most of the time, inside the shell. The male will be territorial now and defend the shell with great vigor. About two weeks after spawning the first fry appear in the opening of the shell. Ad Konings
  • 13. 13 Neolamprologus sexfasciatus occurs in the south- ern half of Lake Tanganyika. It is mainly found, in shallow water, in the intermediate biotope where sand is present between the small heaps of rocks. N. sexfasciatus occurs solitary or in pairs. Breed- ing pairs normally have youngsters which they de- fend until they produce the next spawn. Sometimes juveniles of a size larger than 3 cm are still guarded by the parents. Three different geographical races are known. The holotype was collected on the Zairean coast and probably looked like the fish in the picture above. The first variety that was exported came from Zambia (see photo). It occurs along the entire Zam- bian coast except for the eastern part near the Tan- zanian border. The populations along the Tanzanian coast are completely yellow and have been exported several times. The Golden Sexfasciatus, as this race is called, has been successfully bred in captivity but the Blue Sexfasciatus from Zambia never. In behavior (as far as one can judge from obser- vations in the aquarium) there is a difference be- tween these two races too. The Blue Sexfasciatus shows a much more piscivorous appetite than the Golden variant. The former cannot be kept with A wildcaught female Neolamprologus sexfasciatus from an unknown location in Zaire. Neolamprologus sexfasciatus from Zambia. Neolamprologus sexfasciatus (Trewavas & Poll, 1952) fishes smaller than 5 cm! The race from Zaire seems to have a similar behavior to the Golden Sex- fasciatus. It is impossible to keep more than one pair in an aquarium. In very large tanks (more than 750 liter) two or three females might do well, but usually they quarrel among each other. N. sexfasciatus is a cave-brooder and we there- fore have to supply the pair with some caves. The best way to proceed is to let the female adapt to her new environment first. When she is acclimatized then the male can be introduced. Ad Konings
  • 14. 14 Neolamprologus leloupi was described by Poll (in 1948) from a single specimen which was collected a little north of Moba, on the Zairean coast. Thirty years later, the same author described Neolampro- logus caudopunctatus, a cichlid from the Zambian coast. This fish closely matches N. leloupi in ana- tomical features but lacks the distinct black margin in the tail fin. In 1988 Walter Dieckhoff discovered two other populations of Leloupi-like cichlids on the Tanzanian coast. One of these cannot be distin- guished from the holotype of N. leloupi. The other one closely resembles the Zambian N. caudo- punctatus. Pierre Brichard (1989) concludes in his monu- mental book that N. leloupi occurs in different geo- graphical populations and that N. caudopunctatus should be regarded as a synonym for N. leloupi. Recently Janicki imported another variant from the shores of Zaire. A female of this race is shown in the picture above. The trade name, under which I acquired these pretty fish, is “Red Fin Caudo- punctatus”. I think that Brichard’s opinion will be accepted by most taxonomists, so we should call this little cichlid Neolamprologus leloupi. N. leloupi lives in the intermediate habitat, where rocks lie scattered over a sandy bottom. In Zam- A female Neolamprologus leloupi showing a defensive posture. Neolamprologus leloupi (Poll, 1948) bian waters it is one of the most abundant cichlids in this biotope. Schools of hundreds of individuals are a common sight. In the aquarium, N. leloupi behaves rather peace- fully as long as we keep just one pair. Most of the male’s aggression is directed towards conspecific males. It can be kept with very small fish too. N. leloupi is mature at an age of about one year and may spawn at a size of about 4.5cm. Although it has been observed spawning in an empty Neo- thauma shell, the pair prefers something larger than that. If no cave is present they will start digging one under a stone. During the process the entrance to the cave is concealed by a wall of sand (or gravel). If we provide the pair with a small flowerpot, with its bottom removed and placed upsidedown in the sand, we can chose the position of their territory. They will readily accept the pot and try to conceal it by throwing sand against it. The result is a heap of sand with only the top of the pot sticking out. The size of the territory depends on the size of the fish and the population density of the aquarium, but usually it is about 30 cm in diameter. Any intruder is chased from this area but youngsters (to a size of at least 2.5 cm) from a previous spawn are toler- ated. Ad Konings
  • 15. 15 Neolamprologus mustax is not a common cichlid, either in the wild or in captivity. Its first introduc- tion, in the mid-seventies, was hampered by a wrong identification and by the fact that a fake coloration was applied on a picture of the Cameron Bay vari- ant, the so-called “Blue Mustax”. The first specimens exported to Europe were caught in Cameron Bay. The fish of this population have a grayish-yellow body color and yellow fins. But this race displayed mainly the rather dull gray color when held in dealer’s tanks. Nevertheless, it was named “Blue Petricola”. It was then still an undescribed species but some thought it could be N. petricola. N. mustax is reported only from the southern part of the lake where it has been found in Zambian waters. It is a small, rock dwelling cichlid with a maximum size of approximately 9 cm. Several dif- ferent populations are known. Most of them have fish with yellow fins. Some populations, like those at Cape Chaitika and near Mpulungu, have com- pletely yellow or orange-yellow individuals. The name Mustax means beard which reflects the white coloration on the chin and lower part of the head and which is seen in all known populations. A wildcaught Neolamprologus mustax, collected near Mpulungu, Zambia. Neolamprologus mustax (Poll, 1978) In its habitat N. mustax is difficult to observe as it mainly lives in the dark cracks between the rocks. It is thus also difficult to collect which makes it a rather expensive Tanganyikan cichlid. N. mustax belongs to a group of rock dwelling Lamprologines in which we also find the better known N. leleupi, N. longior and N. cylindricus. All of these species live solitary or in pairs when adult. They are rather intolerant of the presence of conspecifics. It is therefore difficult to accommo- date several individuals in one aquarium. It is a good idea to acquire just one male and a female. If the tank is large enough (about 750 liters with lots of rockwork) both may be introduced at once but it is better to let the female acclimatize first. If not properly fed or kept, N. mustax may lose its deep yellow-orange coloration. It needs a lot of carotene-rich food like Cyclops, Mysis or other al- gae-eating crustaceans. Besides a lot of shelter in the rockwork it needs clean water with a high pH, preferably above 8.0. The colors of N. mustax may fade initially but after treating it right, it will dis- play the golden yellow as brightly as in the wild. Ad Konings
  • 16. 16 Greenwoodochromis christyi was observed under water for the first time in 1988. Walter Dieckhoff found a few individuals in the Zambian part of the lake. He collected a few specimens of which only a female survived transportation back to Europe. One year later some christyi were exported by Chris Blighnaut in Mpulungu. Greenwoodochromis christyi belongs to a group of Tanganyikan cichlids which have a coloration pattern consisting of a few horizontal rows of iri- descent scales. A well-known cichlid from this group is Limnochromis auritus. Previously, all these cichlids were placed in the genus Limnochromis. The species involved are L. auritus, L. abeelei, L. staneri, Greenwoodochromis christyi, G. bellcrossi and Gnathochromis permaxillaris (see next page). The difference between the species in Green- woodochromis and those that remain in Limno- chromis is that the former have more than 48 scales in an horizontal row between the gill-cover and the tail. The species in Limnochromis have a maximum 40 scales in a row. In view of the recent revision of the Malawi cichlids, where the genera are distin- guished mainly by their basic coloration pattern, it A female Greenwoodochromis christyi, collected in Zambian waters. Greenwoodochromis christyi (Trewavas, 1953) would not surprise me if all these species were placed back in their original genus in future. The two species currently in Greenwoodochromis are distinguished from each other by the size of the eye and the shape of the mouth. G. christyi has a relatively smaller eye than G. bellcrossi. The latter also has a steeply inclined mouth, while that of G. christyi is only moderately inclined. Greenwoodochromis christyi has been observed in the intermediate habitat at somewhat deeper lev- els. René Krüter found most individuals at a depth of 25 meters and deeper. Sometimes a pair was ob- served while guarding their offspring. Spawning has not been observed but it is probably similar to that of L. auritus. This means that G. christyi is a mouthbrooder where both female and male take care of their offspring. Greenwoodochromis christyi has proven to be a rather difficult cichlid to keep in an aquarium. A single specimen can be kept, without problems, with other species of similar size but a “pair” need a very large tank. All their aggression is directed towards conspecifics. The solution to this problem could be to let the female adapt to the tank first. Ad Konings
  • 17. 17 One of the most remarkable cichlids of Lake Tan- ganyika is Gnathochromis permaxillaris. Its most interesting features are the very large mouth which opens in a peculiar way (see photo on the backcover) and the shape of the upper lip. It is a rather large cichlid which is known to grow to a size over 18’cm. Gnathochromis permaxillaris lives over muddy substrates and is usually found deeper than 40 me- ters. Although the enormous mouth may suggest otherwise, mostly very small organisms constitute the staple food of G. permaxillaris. These tiny in- vertebrates live close to the bottom, sometimes in the upper layer of the mud. The wide gape of permaxillaris spreads the inflow of water, going into the mouth when the gill covers are extended, over a relatively large area. This weakens the flow so that only small and light particles are carried in. The large cichlid has to “vacuum clean” the muddy bot- tom continuously in order to obtain enough mate- rial to live on. Gnathochromis permaxillaris is probably the most interesting cichlid from Lake Tanganyika. Gnathochromis permaxillaris (David, 1936) The same feeding technique is observed in the aquarium where G. permaxillaris feeds predomi- nantly from the bottom. It takes a while before it adapts to picking up food in mid-water. Gnathochromis permaxillaris is now regularly exported from Zambia albeit in very small numbers. It seems to be a very fragile fish to collect but when acclimatized to the aquarium it proves to be a hardy and undemanding fish. A very welcome factor – completely in contrast to Greenwoodochromis christyi – is its remarkably peaceful attitude towards conspecifics. The sexes can be identified by looking at the ven- tral apertures; the genital opening of the female is noticeably larger than that of the male. Unfortu- nately, there are usually not enough specimens to compare and two males or two females might hide in the same cave without showing any hostility to- wards each other. Ad Konings
  • 18. 18 The cichlids of the genus Trematocara belong to the deepest living fishes of Lake Tanganyika. Representa- tives of this genus were frequently captured at a depth of 200 meters. It is assumed that these cichlids mi- grate freely between the very deep layers and the sur- face layers of the lake. Especially at night they move to shallower water. This means that a special construc- tion of the swimbladder allows the fish to compen- sate for the enormous difference in pressure. During the daytime Trematocara lives in the depths probably together with its food. It is known that a small crusta- cean, a kind of shrimp, undergoes a diurnal vertical migration as well. Masses of this shrimp are found in shallow water at night. Evolution has permitted Trematocara nigrifrons to follow these crustaceans during their daily migrations. This means that these cichlids live in permanent dark- ness. They have well developed adaptations to this circumstance; T. nigrifrons possesses a very large eye and an extremely sensitive lateral organ on the head with which it can register minute movements in the water. The sensory pore system, which is present on the head of other cichlids as well, is greatly enlarged on the head of Trematocara. The organ, which con- sists of pressure sensitive tubules, is visible as small “holes” below the eye. In the aquarium, I noticed that A male Trematocara nigrifrons collected in Zambian waters. Trematocara nigrifrons Boulenger, 1906 food is only accepted (greedily) when it moves. Fro- zen food must be stirred by other fishes before T. nigrifrons eats it. I have seen it hovering above the food on the sand, in a manner similar to Aulonocara, and waiting for a movement to occur. Roger Bills, who is a researcher and collecter of Zambian fishes, managed to catch and ship ten speci- mens of T. nigrifrons. Unfortunately the fish proved to be very sensitive and, after one year, only three sur- vived. Surprisingly I succeeded in breeding them. The male is considerably smaller than the female and, when excited, shows black and white edges to the fins and a black throat. Regrettably I did not observe the spawn- ing; I first knew of it when I noticed the female with a mouthful of pale yellow eggs. After three weeks the fry were about 8 mm in size, when the female spat them out because of a stress caused by netting her out of the tank. The approximately 40 fry immediately ate Artemia and now, three months later, they are about 4 cm long and am I the proud owner of a school of T. nigrifrons. T. nigrifrons is not a colorful addition to any aquarium but for a true Tanganyika fancier, who usu- ally looks further than colors alone, it is a very inter- esting and elegant species. René Krüter
  • 19. 19 The specimens depicted in the photograph above are only tentatively identified as Microdontochromis tenuidentatus. The holotype of this species shows, al- beit in preserved condition, a horizontal, mid-lateral row of black spots. In some preserved specimens these spots appear as vertical bars. The cichlids in the pic- ture above show only such pattern when they are ex- cited. On the other hand, their morphology and their typical mouth strongly suggest that these fish are conspecific with M. tenuidentatus. The holotype, together with about 50 other speci- mens, was collected in Zaïre near Vua. The fish in the picture were collected in Zambia in Cameron Bay, which is not far from Vua. Sporadically, Fishes of Burundi has exported some specimens which also matched M. tenuidentatus in coloration even when they were not excited. These were doubtless collected in Burundi waters. It is known that Cameron Bay harbors some varie- ties (or even distinct species) which are not found at any other place in the lake, for example Petrochromis trewavasae, the so-called Tiger Nasutus or the Ndole Bay variety of Xenotilapia ochrogenys. It is therefore possible that the nondescript cichlid, when not excited, is a geographical variant of M. tenuidentatus which is Two wildcaught specimens of Microdontochromis tenuidentatus of unknown sex. Microdontochromis tenuidentatus (Poll, 1951) only to be found in Cameron Bay. Microdontochromis tenuidentatus is a maternal mouthbrooder. It has been bred in captivity by Volker Puttberg, Dinslaken, Germany. He noticed that the eggs were rather large and that a female broods only a few eggs at the time, although about 20 are deposited dur- ing a spawning. The male does not seem to make a nest or defend a territory. The eggs are brooded for about four weeks before the fry are released. During this period the female continues feeding like she has nothing in her mouth not to be swallowed. In the lake M. tenuidentatus lives in large schools in very shallow water. Most of them are found in the vicinity of aquatic weeds. The mouth is quite protrusible and the upper arm of the upper jaw bulges the skin of the snout. The head is laterally compressed, which is an important characteristic of this species. It seems that M. tenuidentatus feeds from the plankton in the open water as well as from invertebrates which live among the weeds. The teeth in the outer jaws are minute and look rather fragile. The lower pharyngeal bone is very thin and slenderly shaped. The teeth on this bone are minute and pointed. The diet of M. tenuidentatus must con- sists of very soft invertebrates and or tiny organisms. Ad Konings
  • 20. 20 Cichlids from the genus Petrochromis are easily recognized by the structure of the mouth. The nu- merous teeth in the large jaws are permanently vis- ible in most species. Petrochromis orthognathus is the only known species in the genus in which the teeth are less numerous and where the teeth are in- visible when the mouth is closed. A cichlid with the trade name “Petrochromis Tricolor” was ex- ported from Zambia. The specimen which I was able to have a closer look at showed just a few rows of teeth in the mouth. It could close the mouth com- pletely, “hiding” all teeth. The coloration pattern closely matches that of P. orthognathus, i.e. a brown upper half of the body and a lighter lower half. Petrochromis polyodon occurs in several differ- ently colored populations of which at least five have been exported. The cichlid we have previously called Petrochromis famula is the polyodon race from Kigoma. Petrochromis polyodon is easily rec- ognized by the short lower jaw and the generally brown coloration of the females. Petrochromis macrognathus also has a short lower jaw but its mouth is underslung. The “Red Texas Petrochromis” is closely related to the “Texas Petrochromis” from the Tanzanian coast, but is collected on the Zairean side of the lake. A Zairean race of Petrochromis polyodon which is named “Kaiser Petrochromis” in the trade. Above: The “Red Texas Petrochromis”; below: Petrochromis, three new variants Ad Konings
  • 21. 21 Many different populations of Tropheus moorii are known and many more will be found in the future. The genus Tropheus consists of at least six species of which one is scientifically undescribed. This spe- cies, named T. sp. “Black” or—previously—T. sp. aff. brichardi, lives in the northern half of the lake, whereas T. moorii is restricted to the southern half. Although both species have been regarded as conspecific by several authors, Walter Dieckhoff discovered a locality where both species live sympatrically (Kibwesa, Tanzania). A wildcaught male Tropheus moorii, probably from Vua, Zaïre. Tropheus moorii from an unknown location in Zaïre. Tropheus moorii Boulenger, 1898 Ad Konings In Zambia and along the lower half of the Zaïrean coast the populations of T. moorii can be split into two different groups. The holotype of T. moorii was collected near Mpulungu and belongs to the group of light-colored races which are further character- ized by a colored region on the center of the body. In Cameron Bay and north of it, populations of the second group are found. This group consists of dark- colored races which also have a colored cheek. There is no location known where individuals of both groups live together which would have indicated that we are dealing with two different species. Although there is a rather large difference, in col- oration, between the two groups (see photographs), they probably belong to the same species. The two groups therefore represent two main geographical variants. The Kasaba Bay in Zambia might have separated these two groups for a long period of time and they might thus have developed, independently from each other, into the present races. The north- ern borderline between the two groups is not known but a dark-colored variant has allegedly been col- lected near Vua in Zaïre. Further north into Zaïre races of the light-colored group have been found. One of them is shown in the smaller picture.
  • 22. 22 Cyathopharynx furcifer is a rather common and spectacularly colored cichlid from Lake Tangan- yika. Males attain a total length of about 18 cm maximum; females, usually, are considerably smaller. Cyathopharynx furcifer lives mainly over sandy areas in shallow water. However, such sandy bays must be close to rocky shores. The reason why they prefer the rocks nearby is unknown because they predominantly feed from the sandy substrate. The food consists of diatoms which are collected from the upper layer of the sediment on the sand or on the rocks. Large quantities of material are scooped up and processed through the long intesti- nal tract. While feeding, the faeces can be seen growing from the vent. A lot of the material swal- lowed is indigestible. This results in an easy pas- sage of the food through the guts. We must keep this fact in mind when maintaining this cichlid in the aquarium. Cyathopharynx furcifer needs a lot of food but only a little meat. In the lake, males congregate into large breeding colonies. Each male makes its own nest which is a kind of sand-castle with a diameter of approximately 50 cm. The nests are about 30 cm high. The differ- ent territories can be just two meters apart. Breed- This race of Cyathopharynx furcifer lives on the Ruziba reef in the northeastern part of the lake. Cyathopharynx furcifer (Boulenger,1898) ing colonies of C. furcifer are a spectacular sight; sometimes more than 100 colorful males are found together defending their nests. The females remain in large schools, normally close to the rocky coast. Because C. furcifer seems to be dependent on areas with rocks, which are not present along the entire coast uninterruptedly, it has developed sev- eral geographical races. The Ruziba race, which is shown in the photo, was exported from Burundi a few years back. When C. furcifer is collected the fishermen catch the most beautiful males. These males keep their colors for just a few weeks. Rarely do they regain their full palette of colors in captivity, even though they might spawn and produce offspring. Tankraised males, however, show their best colors when still small and usually keep them until something large or rough disturbs them in the aquarium. In the aquarium they need a large area with plain sand where they can build their nest. Only in very large tanks (over 750 liters) can we perhaps have more than one male in color. The presence of a second, smaller, male usually results in the most dominant male retaining his colors. Ad Konings
  • 23. 23 Xenotilapia flavipinnis is a common, sanddwelling cichlid which is found all round the lake. It is found predominantly over sand at depths between 1 and 30 meters. X. flavipinnis lives in schools. The larg- est schools are found in the very shallow water. Foraging schools may number over a hundred indi- viduals. Like all other members of Xenotilapia it feeds by sifting the sand. The food consists mainly of soft-bodied invertebrates, like insect larvae. It seems that X. flavipinnis has traditional forag- ing and breeding grounds and that it is thus restricted to certain areas. This has probably led to the exist- ence of geographical races. Three of these races have been exported to date. The holotype belongs to the northern race of Magara, Burundi. The yel- low variant from Nyanza Lac, Burundi, is shown in the picture. From Zambia a variant is frequently exported which shows a few rows of yellow scales on the flank. Xenotilapia flavipinnis breeds in colonies but it is not a lek-breeder. Pairs separate from the school – the school splits up into many breeding pairs – and establish a rather large territory. Even though we speak of a breeding colony the pairs are usually more than two meters apart. For a small cichlid – the maximum size of X. flavipinnis is about 10 cm The yellow Xenotilapia flavipinnis from Nyanza Lac. Xenotilapia flavipinnis Poll, 1985 – this is a large territory. In order to avoid territo- rial disputes in the aquarium it is therefore best kept as a pair. Spawning occurs at any site in the territory. The site might change during one spawning. There is no visible nest constructed but it seems that the pair chooses a slight dip in the sand. A few days before the actual spawning, the male courts the female by positioning himself with all fins erect and the buc- cal cavity extended. The female answers in a simi- lar way. Without much introductory behavior the female starts laying the eggs which are fertilized by the male while they are still on the sand. After the female has taken all the eggs in her mouth she stays with the male in the territory. After about 9 days the larvae are spat on the sand, in front of the male who picks them up immediately. The male carries the larvae for another 5 to 6 days before he releases the fry in the territory. The fry are tended by both parents – mostly by the male – for another week or two. A further spawning usually occurs within a month in the same territory. Xenotilapia flavipinnis is a rather peaceful cichlid which is best kept with non-predatory cichlids in an aquarium with fine sand on the bottom. Ad Konings
  • 24. 24 Xenotilapia papilio was described from a popula- tion 40 kilometers south of Moba on the Zairean coast. The two individuals shown on this page very likely belong to the same species and are thus geo- graphical variants. X. papilio is a very small sanddwelling cichlid. Its maximum size lies around 8 cm. This very pretty species is mainly character- ized by the underslung and tiny mouth. The teeth are bicuspid, which excludes them from the genus Asprotilapia which is characterized by tricuspid teeth. However, X. papilio closely resembles A. Several geographical variants of Xenotilapia papilio occur in the southeastern part of Lake Tanganyika. This race is known as the “Sunflower Xenotilapia”. Xenotilapia papilio. This variant is known as “Chituta Bay Xenotilapia”. Xenotilapia papilio Büscher, 1990 leptura and is also found sympatrically with the latter. Xenotilapia papilio lives in shallow water but at some locations it may penetrate into depths of more then 30 meters. In the deeper areas is is commonly found in intermediate habitats, where sand and rocks are both present. In very shallow areas it occurs over pure sand. Large schools have never been observed. Usually pairs or small groups are sighted. Like X. flavipinnis it pairs off and defends a territory. Ob- servations of aquarium populations show that once a pair is formed they stay together for several con- secutive spawns. Xenotilapia papilio is best kept as a pair in a community tank as long as there are no predatory species accompanying them. They re- gard similar looking species, such as X. flavipinnis, as competitors which elicit frequent quarrels. Spawning in X. papilio is preceded by fin-dis- plays by both male and female. One or two days later the female deposits the eggs on a site some- where in the territory. These are fertilized by the male while they are still on the sand. The female initially broods the eggs. The male takes the larvae after about 8 to 10 days. After less than a week the fry are released. Ad Konings
  • 25. 25 Xenotilapia sp. “Katete” looks similar to the recently described X. papilio but definitely is another species. It has a shorter snout, a larger eye, a differently shaped dorsal fin and a shallower caudal peduncle. The maxi- mum length of X. sp. “Katete” lies around 7 cm, a little bit smaller than X. papilio. In contrast to latter species, the dorsal fin of X. sp. “Katete” is entirely orange in color. The spines of the anterior part are longer than those of the rest of the fin. The body has a faint blue hue caused by a few rows of blue-reflecting scales. Personally, I find this the most attractive spe- cies among the small, rock-dwelling Xenotilapia. The village of Katete lies near the border between Zambia and Zaïre. In October 1988 we were looking there for a good spot to collect Cyphotilapia frontosa. While I was diving at 35 to 40 meters, I noticed X. sp. “Katete” but at first glance I thought it was X. flavipinnis. Realizing that that species is not commonly found at these depths I looked closer and recognized it from a picture from Horst Walter Dieckhoff in Ad Konings’ book. Xenotilapiasp.“Katete”inhabitstheintermediatezone between sand and rocks, usually on a steeply sloping part of the coast. C. frontosa and Neolamprologus buescheri were among the other fishes of its habitat, which would Xenotilapia sp. “Katete” lives at the deeper levels of the rocky habitat. Xenotilapia sp. “Katete” be a bad combination for the aquarium. With a lot of effort we could only collect a small number of these fragile cichlids of which just five sur- vived the transport to my tank. They were three fe- males and two males. There are no clear morphologi- cal differences between the sexes. Unfortunately, X. sp. “Katete” is very selective when it has to form pairs. I was unlucky that none of the five specimens really liked each other.As long as there had not been a spawn- ing the pair seemed to be doing well; the other three “Katetes” were then “expelled” from the 1000 liter aquarium. These three were housed in a similar sized tank of my friend Willem Bastinck. Xenotilapia sp. “Katete” is a biparental mouth- brooder, this means that the larvae are transferred to the male after the female has brooded them for about 10 days. The problem with my pair started when the male had taken the larvae in his mouth, because then the female started to demolish the male, who reacted by spitting out the larvae. Fortunately, Willem had more success and was able to save some of the larvae after the male had spat them out. Later, my pair also pro- duced some fry which I was able to save from the fe- male’s aggression. Now, after three years, the aquaristic future of these beautiful cichlids looks a bit brighter. René Krüter
  • 26. 26 Cyprichromis sp. “Leptosoma Jumbo” Cyprichromis sp. “Leptosoma Jumbo”, a morph of the Kampemba race. Cyprichromis sp. “Leptosoma Jumbo”, a yellow-tailed morph from an unknown location in Zaïre. Ad Konings
  • 27. 27 The small cichlids from the open water in Lake Tan- ganyika belong mainly to the genus Cyprichromis. There are only two species described in this genus but at least two others are known. The main differ- ence between these species lies in the color patterns of the males. They have gorgeous colors which are clearly seen in their natural environment. Cypri- chromis is spread over the entire lake but only C. microlepidotus occurs in the northern part, north of Nyanza Lac. From observations in Zambian waters (Walter Dieckhoff; René Krüter, pers. comm.) it is known that several species of Cyprichromis may live in direct contact with each other. Cyprichromis microlepidotus, which has a lake- wide distribution, is characterized by small scales. These are smaller than those from any other known species of the genus. It is also larger than C. leptosoma, which is a small and slenderly built cichlid. The latter species differs from C. sp. “Leptosoma Jumbo” by its shallow body and smaller adult size only. C. sp. “Leptosoma Jumbo” can grow to a size, at least under aquarium conditions, of ap- proximately 12 cm whereas C. leptosoma rarely exceeds a length of 9 cm. The fourth species, C. sp. “Leptosoma Zebra”, is recognized by its yellowish coloration – its lack of conspicuous colors – and by the broad vertical bars which are not permanently shown. C. leptosoma (Blue Flash, Isanga race), C. microlepidotus, (Yellow Zambia race), C. sp. “Leptosoma Jumbo” (Black Fin), C. sp. “Leptosoma Zebra” and Paracyprichromis nigripinnis (Blue Neon) have all been seen together in a small area in Chitutu Bay. C. sp. “Leptosoma Jumbo” is a pleasant aquarium fish which normally shows its full colors. It is im- possible to keep this species in pairs. Males actively defend a territory and need the presence of at least Cyprichromis sp. “Leptosoma Jumbo”, a morph from the Kampemba population. C. sp. “Leptosoma Jumbo”, a male of the yellow-tailed morph from the Zaïrean race. four other males to minimize the damage done to the weaker one. Males behave rather peacefully to- wards the females. In three breeding colonies, that I have kept over several years, I have never found a female killed by an aggressive male. Depending on the number of males in the aquarium, they will stake out their territories about 30 to 60 cm apart. The males stay in mid-water and relate the size of their premises to the distance from neighboring males. Females are continuously courted and attracted to the territory. When a female reacts to the male’s invitation, the male bends its body and all its fins, except the ventrals, away from the female. The ven- tral fins, which have a large yellow spot, are held to the outside, in front of the female’s mouth. In- stinctively, the female snaps at the male’s ventral fins. Since spawning takes place in mid-water, this action might indicate to the female the spot where she should deposit her eggs. The color of the spot in the ventral fin does not exactly match that of the egg. I have also observed males snapping at the pre- sented fins of a more dominant male. When a female is ready to spawn she remains inside the male’s territory. Initially she will snap at the male’s ventrals several times before she starts depositing the eggs. He might discharge his milt before the eggs are laid. At a certain moment the male positions himself over the female, all fins ex- tended, and pushes her gently, with his fully opened mouth, on her head. The female releases one or a few eggs and immediately backs up to retrieve it. The sequence of release and backing up may be re- peated several times before the female is led back, by the male, to the center of the territory. Then fol- low a number of “ventral-snappings” before the fe- male discharges the next batch of eggs. The incu- bation period lasts about three weeks.
  • 28. 28 Of all the new imports of the last years, the “Blue Neon Cyprichromis” is the favorite of most Tan- ganyika cichlidists. It is a small and peaceful cichlid with a gorgeous coloration. The maximum size of Paracyprichromis nigripinnis is about 10 cm for aquarium specimens. Its shape closely resembles that of P. brieni, the other species of the genus. The difference lies mostly in their preferred habitat depth, although P. nigripinnis has a noticably larger eye (relatively seen). The latter is also found in deeper regions of the rocky coast than P. brieni, which lives in the upper 10 meters. The “Blue Neon Cyprichromis” is caught at a depth between 20 and 30 meters. In Chituta Bay it lives in large caves or in close vicinity of large boulders. Paracyprichromis nigripinnis has a lake-wide distribution but not all populations have such an attractive pattern as the race shown in the photo- graph. René Krüter (pers. comm.) found two dif- ferent populations in Chituta Bay. The difference between the two is the color of the dorsal fin, which is more red in the other race. Walter Dieckhoff found a race in Kigoma Bay which does not show the “neon” stripes. Paracyprichromis nigripinnis and P. brieni were previously included in the genus Cyprichromis but Paracyprichromis nigripinnis from Chituta Bay is known as “Blue Neon Cyprichromis”. Paracyprichromis nigripinnis (Boulenger, 1901) because of the difference in the number and arrange- ment of the vertebrae they were placed in a new genus. Although the splitting-up might seem a mat- ter of opinion, P. nigripinnis and P. brieni are prob- ably not at all related to the species currently in Cyprichromis. In my opinion, they just show a re- markable case of parallel evolution, whereby spe- cies of two different ancestral origins have devel- oped into similar looking species. The most important distinction between these two genera, however, lies in their spawning behavior. The eggs of Cyprichromis are fertilized inside the female’s mouth (see page 27) but those of Para- cyprichromis are not. Male P. nigripinnis defend a territory alongside a large rock or inside a cave. Females are not at- tracted to the nest by fin display but are seques- tered from the school and then led to the male’s territory. The female swims head-down, close to the substrate, and discharges the eggs. As the eggs pass by her head they are collected by the female. Mean- while, the male stays above or beside the female and discharges – visible clouds of sperm are occa- sionally seen – its milt continuously. He fans the milt towards the falling eggs which are thus ferti- lized before the female picks them up. Ad Konings
  • 29. 29 Of all the different biotopes Lake Malawi has to offer, the rocky regions have brought forth the great- est variety of colorful aquarium fishes. The much less explored sandy areas may still hide many beau- tiful species. Due to the openness of the sandy habi- tats it is difficult, and sometimes impossible, to collect some specimens. Since on the sand there is no place to hide, almost all species living there have a light, silvery-yellow coloration which camou- flages them. Furthermore, the sanddwellers often live in schools which gives them a protection against Placidochromis phenochilus is a very conspicuous cichlid from the sand. MALAWIAN CICHLIDS The blue sanddwellers predators. In contrast to most sanddwellers, there is a group of conspicuously colored cichlids which usually live solitarily or in very small groups. This group con- sists of at least five different species. These have no ancestor in common and do not belong to the same genus. However, they have an important and peculiar part of their feeding behavior in common (see later). The five conspicuously colored sanddwellers are Cyrtocara moorii, Protomelas annectens, Otophar- Ad Konings
  • 30. 30 Placidochromis electra with a digging Taeniolethrinops preaorbitalis. The distribution of Placidochromis electra is not restricted to Likoma Island. This specimen was pictured near Fort ynx selenurus, Placidochromis electra, and Placidochromis phenochilus. The first three species have a lake-wide distribution. P. electra occurs around Likoma Island and probably all along the eastern shores, down to Fort Maguire. P. pheno- chilus, which seems to be closely related to P. electra, has been observed only in the northwest- ern part of the lake. I found it near Mdoka, Chesese, Chirwa Island, and between Selewa and Kasinda. P. phenochilus is characterized by its white lips and blue body. Even juveniles of 6 cm length have the typical dark blue coloration. Its inclusion in Placidochromis is based on its close resemblance to P. electra. Of the about 25 individuals I have seen at several locations, not a single one showed its basic coloration pattern. The vertical barring, which is faintly present in most individuals, is not a diag- nostic feature since species from several other gen- era show vertical bars – especially courting males – as well. The five species of the group have a particular feeding strategy. They are attracted to stirred-up material and usually follow a large sand-sifting spe- cies like Taeniolethrinops preaorbitalis. The food of most sanddwelling species consists of inverte- brates which live and hide in the sand. There are several ways to obtain these. Lethrinops-like spe- cies dig in the sand and sift it through the gills. Species of the genus Aulonocara detect their prey by “listening to the sand”. Other species blow away the upper layer in order to reveal the hiding inver- tebrates. The five blue sanddwellers let other spe- cies do the work. T. preaorbitalis is a large cichlid and its continuous digging efforts create a lot of stirred-up material. Although it seems that the blue sanddwellers select their food from the clouds of material spilled through the gills of the large sand- sifter, they are actually more interested in search- ing the ploughed sand for exposed invertebrates.
  • 31. 31 A male Protomelas annectens is rarely seen in its full blue breeding coloration. Photo taken near Otter Point. Protomelas annectens is the most frequently seen follower of Taeniolethrinops preaorbitalis (photo taken near Fort
  • 32. 32 Otopharynx selenurus from Senga Bay These crawl back into the sand as soon as possible. The intriguing question is, why are these sanddwelling species so conspicuously colored? Especially P. phenochilus, O. selenurus and to a somewhat lesser extent C. moorii, always show a distinct blue coloration. These three species seem to be mostly dependent on a large, sand-sifting host. The other two species are also found foraging on their own. It is therefore possible that their conspicu- ous colors might warn off other followers – espe- cially other species – from joining their host. In sandy regions, cichlids commonly occur in groups or schools, often of mixed species and it would thus be normal for an individual to join such a group. The amount of food revealed by the ploughing ac- tivities of a large digger may not be sufficient for more than one adult follower. If the follower is highly dependent on the host, it may signal its po- sition to the other species by taking on a territorial coloration. In particular individuals of the same species are wary of approaching an “occupied” host. C. moorii and O. selenurus occur in very shal- low water and are rarely found below depths of 10 meters. P. phenochilus and P. electra are usually observed in water deeper than 15 meters whereas P. annectens is found at any level. The differences not only in the basic melanin patterns but also in breeding behavior point to dif- ferent ancestral origins for these sanddwelling cichlids. The breeding behavior of C. moorii has been observed several times by different aquarists. They all noticed that as soon as the female had de- posited some eggs they were fertilized by the male while they were still on the sand. The same sequence is also employed by P. annectens. Neither of the two species build a nest or any structure to indicate the spawning site. Although I have never observed any of these blue sanddwellers spawning in their natural environment, the courting behavior of the males indicates that spawning can take place at any site. Mouthbrooding and fry-guarding C. moorii and P. annectens have been observed on the open sand. P. electra is kept and bred by many hobbyists. Spawning in the aquarium has revealed that the eggs are fertilized inside the female’s mouth, as is the case in most other Malawian cichlids. P. pheno- chilus has yet not been exported as an aquarium fish. Because P. phenochilus resembles P. electra to a great extent it may have a similar breeding behavior, i.e. that the eggs are fertilized inside the female’s mouth. O. selenurus is infrequently exported as Haplo- chromis Nussae. All specimens are collected in Senga Bay. It has been bred in captivity (Baasch, pers. comm.) but details of the spawning sequence are not known yet. The juveniles show the genus- specific blotch pattern until they have reached a length of approximately 7 cm. O. selenurus is a very attractive and peaceful cichlid and makes a valu- able addition to a Malawi community tank. References ECCLES, D.H. & TREWAVAS, E. (1989) Malawian cichlid fishes. The classification of some Haplochromine genera. Lake Fish Movies, Herten, Germany. DRUMMOND, B. (1976) How I keep.....Haplochromis annectens. Bunt. Bull. (Am. Cichl. Ass.), Dec., pp. 29-30. KOCHER, T.D. & MCKAYE, K.R. (1983) Defense of heterospecific cichlids by Cyrtocara moorii in Lake Malawi, Africa. Copeia (2), pp. 544-547. KONINGS, A. (1989) Malawi cichlids in their natu- ral habitat. Verduijn Cichlids, Rotterdam, Neth- erlands. SEIGARS, D. & BERARDO, T. (1979) Spawning the Deep-Water Haplochromis. TFH, Vol. 27; April, pp 4-12.
  • 33. 33 In May 1989, Walter Dieckhoff and I visited the northern region of Lake Malawi, north of Chilumba for the first time. Our trip took us to the Songwe River which is the border between Malawi and Tan- zania. It took us two days and several dives to con- clude that there are no rocky reefs, coasts, or islets between the border and Ngara. The entire coast, here, consists of sandy and swampy beaches al- though small rocks may lie scattered on the beach and suggest that the area is rocky. At Ngara, the most northerly rocky area in Malawian waters, an intermediate type of habitat supports a number of rock dwelling cichlids includ- ing Aulonocara stuartgranti, the race with the or- ange patch on the body. While diving there for the first time I noticed some elongated sand-colored cichlids which moved in small groups through the habitat. At first glance they seemed to be of the genus Aulonocara, a species of the sand dwelling group. They were rather shy and I could take only two photographs. At that time I did not see any in- dividual with male coloration, which could be a fur- ther indication that it belonged to the sand dwell- ing group of Aulonocara. Of course I was eager to have some specimens for a closer examination and asked Saulos Mwale, our boat leader and an ex- A male Aulonocara auditor patrolling its territory. Aulonocara auditor (Trewavas, 1935) tremely skilled fisherman, to catch a few. Saulos needs only two words to describe the desired fish and he will catch it for you, usually much better colored specimens than the ones you had seen. Un- fortunately, he could not find individuals with breed- ing coloration which plainly meant they were not there. However, two specimens were preserved and they later turned out to be A. auditor. At the next location where we dived, at Mdoka, I found the same species again, but again no breed- ing males. Further south, none of us saw A. auditor again. Gary Kratochvil and I went back to the same places in December 1990, where I found A. auditor in breeding coloration. In contrast to the few indi- viduals I had seen in May 1989, we now observed hundreds of them in large schools. Males in full breeding coloration, however, were rare. Most in- dividuals of this species of Aulonocara were found below 15 meters of depth. The few territorial males were found at about 20 m, at the deepest part of the coast where rocks were still present. The territorial behavior of the males resembles closely that of the so-called Chitande Type Aulono- cara; A. auditor might therefore belong to that group of the genus as well. Males in breeding coloration Ad Konings
  • 34. 34 Aulonocara ethelwynnae is found sympatrically with Aulonocara auditor. Aulonocara auditor occurs in schools over sand near rocks. Aulonocara stuartgranti photographed at Mdoka. “defend” their territories with little effort. Other species and females are tolerated at all times within the boundaries of the territory. Competing males are “chased” by raising the dorsal fin and obstructing their way. Under natural circumstances, I have never observed fights, neither in A. auditor nor in Chitande Type Aulonocara. In the aquarium, however, fights may occur. Territorial males live rather far apart in the wild and remain very static. The territory of a male A. auditor usually lies on the sand between some stones or rocks. They were found only at the edge of the rocky habitat. Females occur in large schools and forage over the sand. Within such schools males with faint breeding colors were observed. These males, sometimes showed intolerance towards each other and “chased” each other from the feeding site. No territories were defended within the schools although the school and its members remained rather static. Other species of Aulonocara were found in the schools together with A. auditor. Besides two dif- ferent sand dwelling species, A. rostratum and A.
  • 35. 35 Aulonocara sp. “Chitande Type Kande”. sp. “Jumbo Blue”, foraging individuals of A. ethelwynnae (Northern or Chitande Aulonocara) were also seen. Ngara and Mdoka are the only two locations where two different species of the Chitande Type Aulonocara group are sympatric. Notwithstanding the observation that A. auditor has a similar behavior and habitat preference, it is morphologi- cally clearly different from A. ethelwynnae. The snout of A. auditor is considerably longer than that of all other known species of the Chitande Type Aulonocara. In all the other species the snout pro- file is rounded whereas it is straight in A. auditor. This might mean that A. auditor is able to poke its snout deeper in the substrate than A. ethelwynnae and thus feed from a different population of inver- tebrates. Personally I think that food is abundant in their environment and that those two species are not competing with each other for food. The struc- ture of the teeth in the jaws looks similar in all Chitande Type Aulonocara; A. auditor is no excep- tion. The four to six rows of teeth stand in a rela- tively wide band anteriorly and become a single row on the sides of the jaw. The flat lower jaw, with its teeth, is somewhat reminiscent of that of several Lethrinops species. A. auditor may be more of a sand dwelling species than any other Chitande Type Aulonocara. Territorial males of the other species in this group are found throughout the year; they don’t seem to have a particular breeding season. One exception might be A. sp. “Chitande Type Kande” breeding males of which were rare in June 1989. In December 1990, this species was abundantly present around Kande Island and many territorial males could be observed. At Kande Island, A. sp. “Chitande Type Kande” shares the habitat with several other species of the genus. Two of them, A. kandeensis and A. steveni, are found in its immediate vicinity. It might thus be possible that A. auditor and A. sp. “Chitande Type Kande” breed mainly at a time when other species of Aulonocara decrease their breeding activities. These two species might be unsuccessful in obtain- ing a territory when the other species are still at the peak of their breeding season. References TREWAVAS, E. (1935) A synopsis of the cichlid fishes of Lake Nyasa. Ann. & Mag. N. Hist. Ser. 10. Vol XVI.
  • 36. 36 Abstract The dark-bellied piscivorous cichlid of Lake Ma- lawi is Tyrannochromis macrostoma (Regan), as it was identified by Lewis et al. (1986: 39), who placed it provisionally in the genus Cyrtocara. The darker lower half of the head and body is usual for adults of this species in the lake, but is masked by the blue colour of breeding males and may be lost in unfavourable circumstances. It is thus rarely seen in captive or pre- served fishes. The name T. nigriventer Eccles belongs to a differ- ent species, related to Nimbochromis fuscotaeniatus. The names T. maculiceps (Ahl) and T. polyodon (Trewavas) may be synonyms of T. macrostoma but this is not yet proved. Introduction Our friend Ad Konings, to whom we owe so much for his books on the home life of the cichlids of Lakes Tanganyika and Malawi, has contributed to the solution of the problem of the specific identity of the predator known as “Dark-belly” or “Schwarz- bauch”. But in his article (Cichlidae (BCA) Vol. 11, no. 3) he has made some conflicting statements that A beautiful male Tyrannochromis macrostoma in breeding coloration. Photo by Stuart Grant. The genus Tyrannochromis Eccles & Trewavas, 1989 Dr. Ethelwynn Trewavas need clarification. Konings writes (Cichlidae 11 (3): p. 73) “The so- called Black-belly is thus correctly named T. macrostoma”. But two paragraphs later “the species currently known as T. macrostoma should be correctly named T. nigriventer”. These are apparently mutually contradictory state- ments. In two publications that may be considered “current”, namely Lewis et al. (1986; p. 39) and Ec- cles & Trewavas (1989, p. 97. fig. 43), the name macrostoma is correctly used for the species to which Regan gave that name in 1922. Konings now tells me that by “currently” he meant “currently in the aquaristic trade”.Also in Konings’books, 1989, 1990, macrostoma has been incorrectly used for nigriventer. This is important for us to know because all the eco- logical information we have about T. nigriventer is found in Konings’books under the name macrostoma. While the true macrostoma is there treated under the name T. maculiceps, which may be a synonym of T. macrostoma, or at least the name of a closely related species or subspecies. To explain how this confusion came about let me relate the following, which also shows the value of cooperation between field naturalists, aquarists and taxonomists.
  • 37. 37 A male Tyrannochromis nigriventer in breeding coloration. History of the identification of “Dark-belly” The type species of the genus Tyrannochromis is T. macrostoma, first described by Regan (1922) in the genus Haplochromis on the basis of one specimen of 22.5 cm standard length (26 cm total length). Regan’s figure is reproduced as fig. 43 in our book (Eccles & Trewavas, 1989). The Christy collection contained five other specimens of 12-20’cm total length. All these have white flanks below the mid-lateral black stripe and the markings above this stripe stand out on a pale background. Dr. Digby Lewis and his collaborators (1986; p. 39) have a striking photograph of a fish la- belled Cyrtocara macrostoma with all the lower part of the flank black or dark brown, and their text re- marks on this peculiarity. Could this be the same species as our white-bellied museum specimens? We had verbal reports of the abil- ity of this predator to change colour, but with no de- tails about the conditions in which this might occur. Then, when our book was nearly finished, Eccles found in the Museum a specimen that had been wrongly catalogued as H. polyodon. Its lower flanks were tinged brown. Here, he thought, is our “Dark- belly”, and as it was not either of the named species of Tyrannochromis he described it as a “new species” with the name T. nigriventer, which means “black- belly”. The photograph of this fish is fig. 47 of our book (Eccles & Trewavas, 1989). The chief measur- able difference between this fish and the other species of Tyrannochromis is the short upper arm of the upper jaw (premaxillary pedicel). In T. nigriventer this is less than one third of the length of the head. In the other species it is very long, more than one third of the length of the head, enabling the fish to protrude its mouth as a long funnel without dislocating its jaws. The photograph on p. 6 of Konings (1990a) shows the mouth protruded, but not to its full extent. (This fish is a male in which the overall blue colour masks the adult pattern, of which the only indication consists of a black mark on each scale, more prominent as the ventral surface is approached, giving the flanks a soiled appearance). The paragraphs on p. 103 of Eccles & Trewavas on coloration and ecology apply to T. macrostoma and not to T. nigriventer. So were Dr. Digby Lewis and his collaborators mis- taken in identifying “Dark-belly” with T. macrostoma? This question was answered by our diving naturalists. First,Andreas Spreinat of Göttingen caught and pre- served some “Dark-bellies” and also some other preda- tors. These he sent to the BMNH for our opinions.
  • 38. 38 A juvenile Tyrannochromis nigriventer photographed at Chewere. The tips of the premaxillary pedicels bulge the skin clearly before the eyes. (Size approx. 10 cm TL) Our book was then in the last stages, but after it was published Gordon Howes made some measurements for me. The pre-maxillary test showed that those la- belled “Dark-belly” by Spreinat agreed with T. macrostoma. The others were a good match for T. nigriventer. Spreinat also found that in his photographs the ends of the premaxillary pedicels could be seen bulging the skin. His measurements of these, though naturally inexact, gave a similar result, “Dark-belly” having pedicels whose length went 2.0 to 2.6 times in the head length, whereas those of the other species, identified as N. fuscotaeniatus, went 3.5 to 3.75 times. The relationship between N. fuscotaeniatus and T. nigriventer will be mentioned below. In November 1989 Ad Konings made a further visit to Lake Malawi and he too caught a number of “Dark- belly” and some of a species agreeing with T. nigriventer. The premaxillary test again pointed to T. macrostoma as the correct name for “Dark-belly”. Both divers confirmed that the dark belly is the usual live colour of adult, confident T. macrostoma, in the lake (except blue breeding males). Konings (1989; p. 129) has a photograph of a guarding female that is very dark except on the dorsal side, where the dorsal and dorso-lateral series of markings stand out clearly on a pale background. This photograph is captioned T. maculiceps but Konings now believes, probably cor- rectly, that it is T. macrostoma. The adoption of black colouring when guarding young is reminiscent of the colour changes in the mbuna Melanochromis auratus and may well be a parallel phenomenon. Konings states (1990a; p. 179) that the dark colour is not al- ways present in individuals transferred to a tank. Spreinat saw pale-bellied individuals that were “in- disposed” (letter of 24-6-90). It is understandable that a fish being caught for the pot or for preservation in a museum is not merely “indisposed” but terrified. Hence our pale-bellied museum specimens. The problem of the specific identify of “Dark-belly” is now therefore solved. Konings touched on two other problems in the ge- nus Tyrannochromis. The first is the validity of the two other nominal species, T. polyodon (Trewavas, 1935) and T. maculiceps (Ahl, 1927). Each of these was based on a single specimen. To synonymize T. polyodon it is necessary to show that the greater depth of its body comes within the range of variation of T. macrostoma. To synonymize T. maculiceps it must be shown that its narrow head and interorbital width, as well as possibly its wider band of teeth at any given size, all come within the range of a reasonable sam- ple of T. macrostoma from the southern part of the lake. The type specimen of T. maculiceps was caught at the extreme northeastern part of the lake, a small fish of 163 mm standard length. Other specimens as- signed to this species are from Chilumba and Nkhata Bay. It may be that this is a species or subspecies com- mon in the north. Until these tests have been applied the validity of the names remains an open question. The second problem is the relationship between T. nigriventer and Nimbochromis fuscotaeniatus. They resemble each other in proportions and dentition, but N. fuscotaeniatus has a pigment-pattern which is usu- ally much less regular than in the figured holotype (Regan, 1922 and Eccles & Trewavas, 1989; fig. 79), forming irregular blotches as in other species of Nimbochromis. No such tendency to form blotches is seen in the holotype of T. nigriventer, in which the ventero-lateral brown patch is broader and continu- ous. Konings (1989; p. 222) remarks on the close re- semblance between these two species. Konings does not like to include fuscotaeniatus in A juvenile Tyrannochromis macrostoma photographed at Chitande Island. This specimen is identified by the fact that the tips of the premaxillary pedicels bulge the skin between the eyes. (Size approx. 8 cm TL.)
  • 39. 39 A female Tyrannochromis nigriventer defending her offspring. The photo was taken at Chinyankwazi Island. It shows that a guarding female T. nigriventer can adopt a rather dark coloration on the belly. This female was collected for positive identification. Nimbochromis because of an alleged difference in breeding habits (Konings, 1989; p. 128). The eggs of N. livingstonii are fertilized on the substrate, then taken into the mouth (Konings, 1989; p. 219), whereas those of fuscotaeniatus are said to be fertilized in the mater- nal mouth. The reference for the latter statement is a paper by Dorenstouter (1982), but the identity of the fish in the photograph accompanying this paper is doubtful and the statement requires confirmation. There are other differences between N. fuscotaeniatus and typical species of Nimbochromis and the inclu- sion of fuscotaeniatus requires further consideration. References DORENSTOUTER, C.F., (1982) Haplochromis fusco- taeniatus Regan, 1921. NVC Periodiek (Dutch Cichlid Ass.) April: 43. ECCLES, D.H. & E. TREWAVAS, (1989) Malawian cichlid fishes. The classification of some haplochromine genera. Lake Fish Movies. Herten, West Germany. 335 pp. KONINGS, A., (1989) Malawi cichlids in their natural habitat. Verduijn Cichlids, Netherlands. 303 pp. Illustr. in colour. KONINGS, A., (1990a) Cichlids and all the other fishes of Lake Malawi. TFH Publications, Inc. Neptune City, New Jersey. 495 pp. Many illustr. in colour. KONINGS, A. (1990b) The Malawi genus Tyran- nochromis. Cichlidae (Brit. Cichl. Ass.) Vol 11 (3), pp. 71-73. LEWIS, D., P. REINTHAL, P. & J. TRENDALL (1986) Aguide to the fishes of Lake Malawi national park. World Wildlife Fund. Gland, Switzerland. 71 pp. illustr. in colour. SPREINAT,A. (1990) Zur Revision der Haplochrominen des Malawisees nach Eccles & Trewavas. DATZ (Aquar. Terrar. Z.) (4); pp 245-248, illustr. This article originally appeared in the January 1991 issue of Cichlidae, the periodical of the British Cichlid Association. Details of the BCA may be obtained from page 98.
  • 40. 40 Maravichromis (C.) liemi was previously placed in Caprichromis Eccles & Trewavas (1989). The reason was that it had a strongly inclined mouth, a broad lower jaw and a specialized feeding behavior. Although it is functional to split large genera into a few smaller ones, monotypic or oligotypic genera should be avoided when possible. The peculiar structure of the mouth of liemi and orthognathus is related to the feeding behavior but other species in the lake also have simi- lar mouth structures and comparable behavior. In the genus Protomelas there are several, undescribed spe- cies (e.g P. sp. “Paedophage”) with a similar mouth structure, but none of these or other species like Diplotaxodon greenwoodi, have been put or will be put into a new genus or into Caprichromis. Personally I think the different mouth structure is insufficient justi- fication for erecting a new genus for these two species. M. (C.) liemi is reported to steal eggs and larvae from mouthbrooding females (McKaye & Kocher, Animal Behav. 31, 1983), specifically from P. pleurotaenia. The authors probably meant Nyassachromis microcephalus as P. pleurotaenia remains close to the bottom and mouthbrooding females are not found in schools. I have observed M. (C.) liemi at many different lo- cations all round the lake and it seems likely that their diet is not restricted to larvae and eggs alone. I have A male Maravichromis (Caprichromis) liemi in full breeding coloration photographed at Eccles Reef. A subadult Maravichromis (Caprichromis) liemi. Maravichromis (Caprichromis) liemi (McKaye & McKenzie, 1982) observed several times that M. (C.) liemi attacked male cichlids as well, preferably those with parasites (Argulus) on the throat region. It is therefore possible that M. (C.) liemi feeds also, or even primarily, on these parasitic invertebrates since fish with Argulus on their throat are very common. M. (C.) liemi is normally found near rocks whereas M. (C.) orthognathus prefers the open sand. Breeding males congregate into colonies and construct a sand- castle nest (more than 1 m in diameter) against a rock. Mouthbrooding females stay in schools and probably release their fry in the rocky habitat. Ad Konings
  • 41. 41 Sciaenochromis gracilis was described from three specimens in the Christy collection. The total length of the largest specimen measured 217 mm. Some sin- gle specimens have been imported alive under trade names such as “Torpedo” or “Dark Line Torpedo”. Recently, the export of S. gracilis has taken a more frequent character. This is certainly caused by the ef- forts of Stuart Grant to comply with the increased de- mand. The distribution of this rare species seems to be re- stricted to the southern part of the lake. Observations about their feeding and breeding behavior and about their breeding coloration are not published yet. The specific name (gracilis = slender) is aptly cho- sen. The narrowly pointed head with a slightly down- ward curved profile is a striking feature that reminds one of Nimbochromis linni.Aband, consisting of small spots, runs diagonally from the nape to the upper part of the tail (peduncle). In this respect it could be mis- taken for S. spilostichus or Maravichromis formosus. Maybe gracilis should be placed into Maravichromis as has been suggested before (Konings, 1989). Its large mouth, its teeth and its shape presume a predatory lifestyle. The strong tail indicates a fast swimmer with ability to spurt. A courting male Sciaenochromis gracilis photographed in the author’s aquarium. Sciaenochromis gracilis (Trewavas, 1935) Outside the breeding period S. gracilis is a lively swimmer. Every now and then they suddenly stop and move their eyes noticeably. Although S. gracilis be- haves rather peacefully we should not house them in too small quarters. Their behavior changes during the breeding period. Then S. gracilis is able to spurt over large distances without apparent effort. Courting is rather impetuous and intensive, but the female is not hurt. Males defend a territory and con- struct (in my tank) a crater nest with a diameter of about 50 cm. The eggs are very small and are ferti- lized inside the female’s mouth. Since the eggs are small many fry can be expected. Females with a size of approximately 15 cm may release over 100 fry. Ju- veniles are, in comparison to other predators of simi- lar size, very small and slim. They grow rather slowly, probably because they show the same behavior as the adults, i.e. they frequently stop moving and search the area with their eyes only. Food must then be brought into their mouths to let them eat! At a size of about 2- 3 cm they defend small feeding territories and quarrel among themselves. S. gracilis grows to a maximum size of about 25’cm and is in shape, coloration, and behavior a valuable addition to any large aquarium. Peter Baasch
  • 42. 42 Otopharynx decorus is a small to medium sized cichlid which is found predominantly over the open sandy bot- toms of the lake. Its distribution encompasses probably the entire lake because specimens have been collected at Vua as well as in the southern part of the lake. The maxi- mum size of O. decorus is about 18 cm (total length); females are only a little bit smaller than males. This sanddwelling species lives in small groups, usually numbering not more than 6 members. Solitary individuals are frequently observed, too. In Decem- ber, when I made the observations, there was some- times one male individual in the group that showed some of its breeding coloration. Territorial males have not been seen. Members of the genus Otopharynx usu- ally spawn near rocks. It is therefore possible that O. decorus also spawns close to a rocky coast and that the males which I was able to observe were foraging and not breeding. The feeding behavior of O. decorus is rather char- acteristic of an insectivorous cichlid. With its large eyes it carefully screens the sand. Only after it has located something interesting does it bite into the substrate. Its food consists mainly of invertebrates which hide in the sand. The teeth on the lower pharyngeal bone are somewhat enlarged in the center, which is a common A male Otopharynx decorus taken at a depth of approximately 18 meters in Senga Bay. Otopharynx decorus (Trewavas, 1935) feature in cichlids which live on crustaceans and in- sects. The feeding behavior closely resembles that of Maravichromis sp. “Double Spot” (see next page). Both species have a small mouth with which they se- lectively pick their food from the substrate. Both spe- cies are found sympatrically. The neutral coloration pattern consists of three rather large blotches on the flank. These blotches are not al- ways distinct. When observed in the somewhat sub- dued light at a depth of 20 meters, the spots are almost invisible. I have never seen them black as in O. (Ctenopharynx) nitidus with which it is commonly found sympatrically. The coloration of the body of O. decorus is like that of the sand – light beige. Among the sanddwelling cichlids of the lake it is rather easily recognized by the combination of the small mouth and the size of the fish. When observed in its natural environment, its behavior will immedi- ately allow an identification. Identification of aquarium or preserved specimens is simplified by the fact that O. decorus has a high number of soft rays in the dorsal fin, 13 or 14, whereas most other Haplochromines have less than 12. It has, furthermore, a high number of scales on the flanks, higher than any other known Oto- pharynx. Ad Konings
  • 43. 43 Maravichromis sp. “Double Spot” is a rather small, sand-dwelling cichlid. Its maximum size is known to be less than 14 cm (total length). It has a rather wide distribution since it has been found on the east coast near Fort Maguire, as well as near Mdoka in the north- ern part of the lake, and near Chintheche. It has, how- ever, never been seen in Senga Bay, although cichlids are frequently collected there. M. sp. “Double Spot” is characterized by a diago- nal row of three double-spots. In some specimens the spots are so large that they form a broad, almost solid black band. This feature helped me decide to put it in the genus Maravichromis and not in Otopharynx. The “Double Spot” prefers the open sandy regions and is usually found at between 5 and 30 meters depth. Most individuals forage on their own, rarely two or three individuals are seen together. The coast near Fort Maguire is inhabited by a rather dense population, al- though they never forage in groups or schools. At the two other locations where I have seen M. sp. “Double Spot”, only single individuals were found. The fish of the east coast population have a yellow coloration on the body and especially on the fins, whereas speci- mens from the other locations are more silvery and have colorless fins. Maravichromis sp. “Double Spot” scrutinizes the sand for something edible (Fort Maguire). Maravichromis sp. “Double Spot” The mouth of M. sp. “Double Spot” is very small and is utilized to pick small invertebrates from the sand. The substrate is therefore scrutinized for anything ed- ible. The characteristic posture of this cichlid is poised over the sand, visually screening the area in front of the head with its large, movable eyes. The fish swims from one place to the next and everywhere it halts and scrutinizes the sand. In this respect M. sp. “Double Spot” resembles Otopharynx decorus. The latter spe- cies also forages in small groups. The specialized feeding behavior of M. sp. “Dou- ble Spot” might also be used for screening parasites and fungus on the body and fins of sick fish. Once I observed a female Nyassachromis prostoma, with a visible fungus in the dorsal fin, who presented the ail- ing fin by laying itself on the sand in front of a forag- ing “Double Spot”. At the same moment a M. labidodon disturbed the scene, so I could not see whether M. sp. “Double Spot” would actually clean the prostoma’s fin or not. It is, however, likely that it does, otherwise the ailing fish would not present its fin in the way it did. The specific coloration pattern of the “Double Spot” is unique among the haplo- chromines of the lake and might function as a signal to other species that it is a “Cleaner”. Ad Konings