Eggspots Elsewhere - Welt der Fische / World of Fishes

In search of haplochromines in Egypt - 1. Ismailia 

A little known Pseudocrenilabrus species from East Africa 

Maylandia or Metriaclima – the case for Metriaclima

eggspots 

The journal on fishes with eggspots 

Number 4 - Date 19.10.2010 

Pages 1-48 

Editor 

ERWIN SCHRAML, Haferstraße 18c, D-86179 Augsburg 

Tel.: +49 821 86 886, Fax: +49 821 86594, E-mail: schraml.e@web.de 

Consultant Editors 

MARY BAILEY BA (freelance aquatic consultant/translator) 

general cichlids/aquarium maintenance; preparation of English text. 

Dr PETER BURGESS (consultant to Aquarian) - fish health. 

MARTIN GEERTS (author and taxonomic specialist) - taxonomy and systematics. 

Dr ULRICH K. SCHLIEWEN (ichthyologist) - genetics and Congo. 

CIP-Titelaufnahme der Deutschen Bibliothek 

Eggspots (English ed.) : The journal on fishes with eggspots. - Augsburg : SCHRAML 

ISSN 1869-2362 

Copyright © 2010 by ERWIN SCHRAML, Augsburg, Germany 

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 or otherwise, without 

the prior permission of the copyright owner.

Editorial 

In this issue of eggspots we not only continue our report 

on a trip to Egypt, but also present for the first time, in 

words and pictures, a Haplochromis species caught there 

and compare it with other North African species. 

Additional articles deal with new information on the 

phylogeny of cichlids in general, the description of a new 

Haplochromis species, the discussion of a taxonomic problem (which luckily 

doesn’t affect us greatly), and the sad demise of an excellent cichlid 

association. We also devote several pages to a little-known 

Pseudocrenilabrus species from Tanzania, and finally we have a response to 

the Maylandia article in a previous issue of eggspots, in which the 

arguments of the Metriaclima supporters are explained in detail. The issue 

is completed by Eggspots Elsewhere and a short book review. 

This issue also represents an experiment, in that we are allowing readers to 

simply download the journal from the Internet with payment on a voluntary 

basis. I would therefore like to take this opportunity to request all readers 

to make a contribution to the not inconsiderable costs involved in producing 

each issue. You will find details on the final page. 

In this regard, I hope you will read this fourth issue with great interest! 

Erwin Schraml 

eggspots No. 4 3

Contents 

Impressum ............................................................................................ 

Editorial ................................................................................................ 

Articles: 

In search of haplochromines in Egypt - 1. Ismailia................................ 

A comparison of Haplochromis sp. “Ismailia” with other North African 

species.................................................................................................. 

Cichlids (and other acanthopterygian fishes) phylogenetically 

reclassified ........................................................................................... 

A new Haplochromis described from the Katonga................................ 

The gender of the genus Haplochromis and other –chromis............... 

The end of an era.................................................................................. 

A little known Pseudocrenilabrus species from East Africa................... 

Maylandia or Metriaclima – the case for Metriaclima............................ 

Eggspots Elsewhere ............................................................................. 

Cover photo: Haplochromis sp. “Ismailia“ 

Page 

2 

3 

5 

12 

25 

27 

29 

31 

32 

39 

46

The story behind my trip to Egypt with ANDREAS DUNZ has already appeared in an 

article in the previous issue of eggspots (DUNZ & SCHRAML, 2010). Ismailia was the goal 

of the first stage of our expedition. This town can be reached without problem from 

Cairo in a few hours via a well-constructed road – at least once you have finally 

managed to escape the horrendous traffic chaos of the capital. Ismailia was mainly of 

interest to my travelling companion because it is the type locality for two little known 

tilapiine cichlids, Oreochromis and Tilapia ismailiaensis. Both were discovered in the 

Ismailia Canal, which runs for 128 kilometres from Cairo to Ismailia and there enters 

Lake Timsah, which is nowadays connected to the Suez Canal. The precursor of the 

Ismailia Canal was the so-called Bubastis Canal, which was constructed in ancient 

times (610-595 BC). There have even been suggestions (SCHÖRNER 2000) that a canal 

was constructed even earlier, at the time of the Pharaoh Sesostris I (1956 to 1910 

BC). These historical details demonstrate that although the Ismailia Canal is an 

artificial waterway, it has perhaps been in existence as an aquatic habitat for almost 

4000 years and hence it is quite conceivable that its own distinct species may have 

evolved there during this long period of time, even if they are probably closely related 

to species from the Nile. The more so in that the canal has probably periodically 

become silted up with sand in places and hence cut off from the Nile. In addition, 

prior to the construction of the Aswan Dam the Nile inundation probably regularly 

A “side-arm” of the Ismailia Canal, about six kilometres from the town of Ismailia. 

eggspots No. 4 

In search of haplochromines in Egypt 

- 1. Ismailia 

by ERWIN SCHRAML 

5

eached Lake Timsah via the Wadi Tumilat, which is again an additional indication of 

the likelihood of a distinct fauna in the Lake Timsah Tinsah region. 

Andreas had previously measured the type specimens of the two tilapiines from the 

region at the Natural History Museum in London and studied their descriptions. The 

differences from the better-known Nilotic species were not very significant, and so we 

were very interested in 

whether live specimens 

might perhaps exhibit 

greater differences. Unfor - 

tun ately we were unable to 

catch any tilapiine fishes 

ourselves in the entire area 

around, Ismailia, and the 

few that we found in the 

catches of the fishermen 

looked just like normal 

Oreochromis niloti cus. The 

only interesting non-cichlid 

that we saw was an African 

sardine (Chela ethiops bibie), 

which was searching for food at the water’s surface almost at the end of the Ismailia Canal. 

Immediately after the last sluice there were anglers busy catching fishes in the sea 

water. A glance into their catch buckets revealed that in the main they were capturing 

rela tively small wrasses. Were these intended as bait for larger fishes? 

Communication with the anglers, as with other passers-by, was difficult, as the 

majority of the people here, outside the tourist regions, spoke only Arabic, but we did 

not. 

The Ismailia Canal is completely concreted in the area of the town, and mostly with 

very high embankments. The water level lies so far below that it would be difficult to 

try and fish from the bank. Because the bottom was almost entirely bare of cover we 

decided not to bother. So initially the only place where we were successful in catching 

6 

Left: The Ismailia Canal is slightly 

higher than Lake Timasah where it 

enters the lake. Sluice-gates prevent salty 

water from flowing back into the canal, 

whose water is used for the irrigation of 

fields cultivated for agriculture. 

Below: an African sardine (Chelaethiops 

bibie) at the water’s surface, picking out 

food that had landed there. 

SCHRAML: Haplochromis sp. “Ismailia“

Everywhere in Egypt we encountered 

North American faunal elements: 

Gambusia (right: a male G. affinis) 

and crayfishes (below: Procambarus 

clarkii, here a juvenile). 

Bottom: Hopefully at least the shrimps 

that were also to be found almost 

everywhere are “true Egyptians”. 

our surprise, crayfishes as 

well. They may well have 

been another North 

American species, Pro - 

cam barus clarkii, which 

we also frequently encoun - 

tered in waters in Egypt 

that we visited subsequently. 

fishes was a side channel, 

around six kilometres 

outside the town, where 

we captured the North 

American Gambusia that 

we encountered practi - 

cally everywhere in Egypt 

and which had probably 

been introduced a long 

time ago to combat 

mosquitoes. Other bycatches 

included shrimps and, to 

But – to my great pleasure 

in particular – we also 

caught a number of 

haplochromines. The current 

list on FishBase doesn’t include any described species of these cichlids from Egypt at 

all. But in his book on the fishes of the Nile BOULENGER records a species that he terms 

Haplochromis bloyeti, and if I remember correctly this name and that of Haplochromis 

loati were until recently still included on the FishBase list for Egypt. And as already 

mentioned in the earlier article on the Egyptian Mouthbrooder (Pseudocrenilabrus 

multicolor), the literature does, however, contain indications regarding as yet 

undescribed species from Lake Maryut, from the Nile in the area of Luxor, and from 

the Fayoum region. And now also from the vicinity of Ismailia. 


7

The specimen in the upper photo has an unmistakeably longer snout than the one below. Note also the stripe pattern. 

From the same site as the specimen above, a Haplochromis with a rounder head profile. The cuvette photos were taken immediately after 

capture. 

8 


The easiest place to find these fishes was beneath marginal plants trailing in the water or 

among truly submerse vegetation. It turned out that we were able to distinguish two 

different phenotypes among the fishes that we caught, specifically some with a long and 

some with a shorter and rounder snout. Immediately after capture all the specimens we 

caught had a vertical stripe pattern on the sides of the body, but this was hardly visible at 

all later on. In the aquarium they practically always exhibited a horizontal pattern 

consisting of a mid-lateral band and a subdorsal lateral band. The facial mask includes the 

Hemichromis letourneuxi from a subsidiary canal in the vicinity of Ismailia. 

entire spectrum of markings: nostril stripe, interorbital stripe, supraorbital stripe, nape 

band, and lacrymal stripe, and there is even a hint of a preopercular vertical stripe visible. 

While I had the more pleasant job of photographing the specimens we had captured, 

Andreas had to ready for preservation those that we couldn’t take back with us alive. He 

was also responsible for the DNA samples and writing up details of, for example, the 

collecting location. This division of labour meant that we always got things done quickly 

and were able to cope with a good deal of work. 

As already described in the last issue of eggspots, our trip to the Ismailia Canal included a 

visit to the Birket Abu Jumas. On the way back to Cairo we stopped at yet another side-arm 

of a canal which produced additional Haplochromis and Hemichromis letourneuxi; we had 

already caught the latter in the Abu Jumas, but here they were somewhat paler in colour. 

On the return journey we also investigated a place, around 50 kilometres from Ismailia, 

that appears on the satellite map as a lake-like widening of a flowing water. It lay right next 

to the road in the middle of nowhere, and appeared to consist of an artificial arrangement 


9

with several pools. But to our disappointment we discovered that it was no more than some 

kind of effluent disposal system and the artificial pools were perhaps settling beds. 

To sum up this first day: while we might not have found the tilapiine fishes we were looking 

for, instead we had discovered what was probably a new Haplochromis species and a new 

location for Pseudocrenilabrus multicolor and Hemichromis letourneuxi. 

Male Haplochromis sp. “Ismailia” courting a female (in the background). 

Aquarium observations on the Haplochromis from Ismailia: 

Although Haplochromis sp. “Ismailia” is only a comparatively small species, territorial 

males are exceptionally aggressive towards not only rivals but also females that are 

unwilling to spawn. I was, however, able to establish that in winter (I wasn’t able to 

determine the precise months but it was probably at least from December to February) all 

courtship activity came to a halt. During this period females and even males were in 

harmony with one another, without any aggressive encounters at all. The rest of the time 

the male was constantly busy displaying the moment a female came near, or actively 

searching for a female to court. At the peak of courtship arousal all the bars on the sides 

of the male’s body disappear. By contrast, the females are invariably pale grey to creamcoloured 

all over. 

A relatively shallow spawning pit is constructed by the male, who then entices ripe females 

into this depression. Spawning takes place in the manner typical of all haplochromines, in 

a head to tail position with the pair circling. The females brood for only around two weeks 

and harbour up to around 50 fry in their mouths – a relatively small number in comparison 

to other Haplochromis. The young are taken back into the mouth at any disturbance, 

10 


Two females of Haplochromis sp. “Ismailia”. 

During the phase of highest arousal the lateral stripes disappear completely in males of Haplochromis sp. “Ismailia”. 


11

though if the latter is too great then it may happen that the female takes flight before 

collecting up the fry. It is then sensible to separate the mother from the young. The further 

rearing of the fry is problem-free and possible using crumbled flake as food. 

References: 

DUNZ, A. & SCHRAML, E. (2010): Fresh blood from Egypt - Pseudocrenilabrus multicolor imported again after many years. eggspots, 3: 5-16. 

SCHÖRNER, H. (2000): Künstliche Schiffahrtskanäle in der Antike. Der sogenannte antike Suez-Kanal, Skyllis, 3 (1): 28–43. 

A comparison of Haplochromis sp. “Ismailia” with other 

North African species 


To date only two haplochromine species have been described from northern Africa and the 

Levant. One is Haplochromis desfontainesi from Tunisia and Algeria, the type species of the 

genus Astatotilapia, and the other Haplochromis flaviijosephi from the Jordan drainage, 

which is at the same time the only species of the group whose distribution lies outside of 

the African continent. 

When PELLEGRIN (1904) erected the genus Astatotilapia to accommodate African species 

which he saw as standing between the genera Tilapia and Paratilapia, he chose as type 

species Sparus desfontainii 1). He regarded as characteristic inter alia the change from 

bicuspid to unicuspid teeth on attaining adulthood that gives the genus its name (astatos 

= variable). 

GREENWOOD (1979), who revalidated Astatotilapia (after REGAN (1922) had treated it as a 

synonym of Haplochromis), spends two pages describing the characteristics of taxa that he 

assigns to this genus and in addition provides a page-long diagnosis, but his 

characterisation is so broad that no clear picture emerges. He himself even expresses 

doubts (GREENWOOD 1980) as to whether the genus, as defined by him, is monophyletic. 

MEYER et al. (1991) and MEYER (1993) confirm on the basis of molecular studies that several 

lineages have been lumped together with no justification. In addition LIPPITSCH (1993) found 

uniform scalation characters in fluviatile Astatotilapia, but that these differed from those 

of the lacustrine species, for which reason only a distant relationship can be assumed at 

best. 

1) PELLEGRIN assumed, however, that this species was described by LACÉPÈDE as a Labrus and not, as was actually the case, in the genus 

Sparus, although he does also list the correct combination, as it had been used in GERVAIS (1869). Presumably he hadn’t actually read the 

work of LACÉPÈDE and was simply adopting the combinations mentioned in BOULENGER (1899). An indication of this is that both authors 

for the first time use the species name desfontainesi rather than desfontainii, but cite the latter as the original spelling. GERVAIS (1869) 

and SAUVAGE (1877) both previously used desfontainii, and this was undoubtedly known to both BOULENGER and PELLEGRIN. For 

unknown reasons (perhaps only a printer’s error?) we find both spellings in LACÉPÈDE, on p. 54 as Sparus Desfontainii and p. 160 as 

Sparus desfontaines (explicitly cited as the scientific nomen representing the similar-sounding French name). After BOULENGER (1899) 

desfontainesi was predominantly used in the literature, but following the publication of CLOFFA IV (1991) desfontainii resurfaced, 

although GREENWOOD (1979 - in a footnote) had already explained that BOULENGER corrected the name because he assumed that 

LACÉPÈDE wished to use this spelling as the name was chosen to honour a M. Desfontaines. The ICZN allows/requires correction of a 

name only if “32.5.1. If there is in the original publication itself, without recourse to any external source of information, clear evidence 

of an inadvertent error, such as a lapsus calami or a copyist's or printer's error, it must be corrected. Incorrect transliteration or 

latinization, or use of an inappropriate connecting vowel, are not to be considered inadvertent errors.” Because LACÉPÈDE used both 

spellings and in addition states that the species was to be dedicated to “citoyen Desfontaines” (Citizen Desfontaines), who had discovered 

the fish in Tunisia, a spelling or printer’s error appears likely. Hence in the event that if, for formal reasons, BOULENGER (1899) cannot 

be regarded as the First Reviser (article 24.2 of the Code) , I assert that ‘desfontainesi’ should nevertheless be accepted as the correct 

spelling of the species name. 

12 SCHRAML: Comparison of North African species

Threatening male Haplochromis desfontainesi. 

Female Haplochromis desfontainesi. 


13

On the other hand, other authors recognise morphological indicators that speak against 

any splitting into separate genera related to Astatotilapia. Thus WITTE & WITTE-MAAS (1987) 

had previously discovered, from material used by GREENWOOD & GEE (1969) for their 

descriptions, that the transition between Yssichromis and Astatotilapia is fluid. SNOEKS 

(1994) found intermediate species from Lake Kivu such that there was no longer any clear 

separation between Astatotilapia and Gaurochromis. 

All in all, Astatotilapia is an example of why authors such as VAN OIJEN (1996) have 

recommended that all haplochromine species from the Lake Victoria basin should for the 

time being be left in the genus Haplochromis. This recommendation has long since been 

adopted for other haplochromines (with exceptions such as the so-called Malawi 

Haplochromis) by CLOFFA and well-known online databanks such as the Catalog of Fishes 

and FishBase. However, recently the name Astatotilapia has again been used in these 

online databanks. Because, in the absence of a newer and more precise diagnosis for this 

genus, it is difficult to determine what actually constitutes Astatotilapia and what does not, 

I would like to make a personal plea for the continued use of Haplochromis for the time 

being. In the subsequent course of this article I will, nevertheless, use the genus names 

that the authors in question have regarded as correct in the works cited. Even if I had 

decided to do otherwise, a certain amount of confusion in this regard would still have been 

inevitable. 

A recent genetic, albeit only mitochondrial, study by GENNER & HAESLER (2010) finds that 

from a phylogenetic viewpoint A. desfontainesi belongs to the modern haplochromines of 

the mainly East-African/Nilotic lineage. It is thus relatively similar to A. flaviijosephi from 

the Levant, but also to A. burtoni from 

Lake Tanganyika. On this basis it 

belongs to a relatively old and 

apparently monophyletic lineage. 

LIPPITSCH (1990) describes the 

structure of a flank scale in A. burtoni 

(as type 10), a typical Astatotilapia. 

According to her granulation is visible 

over the entire exposed part and 

totally obscures the ring structure. It 

consists of blunt grains or tubercles 

which are often wart-like. Ctenii are 

visible only at the very edge, if at all, 

and are rather small. 

Haplochromis desfontainesi 

Haplochromis desfontainesi is known 

only from Tunisia and Algeria. It is 

regarded as a relict in permanent 

waters in the basin of the former Chott 

Right: Flank scale of H. desfontainesi. As LIPPITSCH 

describes for type 10, the circuli on the exposed part of 

the scale are totally obscured by granulation and ctenii 

are present only at the extreme edge. 

14 

SCHRAML: Comparison of North African species

Details of a flank scale of H. desfontainesi: left the small number of ctenii and right the radii. 


Above: Scale size alters only gradually during the transition from flank to 

breast. 

Left: In some specimens of H. desfontainesi the first scale of the lower lateral 

line (F) is sited anterior to the last scale (L) of the upper lateral line. 

15

Above: The lower-jaw teeth are largely unicuspid in adult males of Haplochromis desfontainesi. The large photo shows the right-hand 

side (front to the right), the small photo two teeth on the left-hand side. 

paleolake and originates from an 

epoch that probably came to an end 

with the last ice age. Back then the 

Sahara was also moister and it is 

known from Stone Age paintings that 

there were formerly also large 

animals there that today are known 

only from the savannas of East 

Africa. In 2006 I myself brought back 

a number of juveniles from Tozeur 

(SCHRAML 2007a, b), but despite an 

exhaustive search was unable to find 

any other sites for the species in 

Tunisia. Hence it is categorised as 

seriously endangered (EN) in the 

IUCN Red List. However, JÖRG FREYHOF (pers. comm.) has subsequently been able to rediscover 

the species in Lala as well. 

Maximum length in males can be up to 15 centimetres. The taxon has a relatively large number 

of lateral-line scales (total for both parts) for a fluviatile species (according to GREENWOOD 

(1979) 31-33 vs. 28-30 in other Astatotilapia; a count of more than 30 is generally found only 

in lacustrine haplochromines (GREENWOOD 1980)). I have established that in specimens of H. 

desfontainesi with a high number of scales in the lateral line the count for the median series 

of scales is lower. In one case the number of pored scales was 32, but that for the median 

series only 29. In many haplochromines the two values are identical. The reasons for the 

difference are on the one hand that the species is high-backed and hence the upward curvature 

results in more scales being included in the upper lateral line, and on the other that the lower 

16 

Anterior teeth of a male H. desfontainesi, in part with unicuspid teeth that look 

as if they were originally bicuspid, but the minor cusp has been lost. 


section doesn’t continue from a point posterior to that at which the upper terminates, as in 

many other haplochromines, but the two parts overlap. 

The size difference between the scales on the breast and flanks is not marked and takes place 

gradually, just as GREENWOOD (1979) also describes as typical for Astatotilapia. As can be seen 

from the photos showing flank scales, H. desfontainesi corresponds precisely to the type 10 of 

LIPPITSCH. GREENWOOD’s statement that in Astatotilapia the ctenii are generally strong and 

extend around the greater part of the scale’s free margin does not apply in the case of the type 

species, where they are rather small and cover only a small part of the scale margin. 

In the large specimens I studied (only males were available) the jaw dentition was 

predominantly unicuspid, although it was possible to discern individual variation. In some 

specimens the teeth are comparatively robust and the crowns rounded and pointed, and 

slightly incurved. In other individuals they are noticeably more slender and look as if they may 

actually be bicuspid teeth whose 

minor cusp has been partially worn 

away. The pharyngeal dentition 

resembles that of H. flaviijosephi, of 

which GREENWOOD (1979) writes that 

it constitutes an exceptional case in 

Astatotilapia, as only this species 

develops somewhat enlarged teeth 

with submolariform crowns in the 

central series. 

The gill rakers correspond to 

expectations for Astatotilapia, 

following GREENWOOD (1979) in form 

First gill arch of H. desfontainesi. 


Left and below: Pharyngeal dentition of H. 

desfontainesi: the large submolariform 

teeth in the upper centre are distinctive and 

typical, and their arrangement is regarded 

as characteristic for the genus Astatotilapia. 

17

Male H. flaviijosephi in courtship coloration. 

Ripe female H. flaviijosephi. 

18 


Above: First gill arch of H. flaviijosephi with large gill rakers. 

Right: Flank scale of H. flaviijosephi. The short but broad radii 

(so-called secondary radii) are striking. 

Below: Magnified details of (left) the free edge of the flank scale 

with its relatively small ctenii, and (right) the part normally 

covered by the adjoining scale, showing the radii. 


19

20 


Above: The teeth have been exposed in this preserved specimen of H. flaviijosephi (a male measuring 71.1 mm SL). Some of the teeth 

appear almost unicuspid, though this appears to be attributable more to reduction (wear?) of the minor cusp than to their being true 

unicuspid teeth. 

Photos page 20: Teeth of a 55.7 mm SL male of H. flaviijosephi: lower pharyngeal dentition (top); jaw teeth in anterior (centre) and lefthand 

lateral (bottom) view. 

and number (eight rakers, some slender and moderately long, some shorter and blunt). It is 

noteworthy, however, that there is a rather delicate, well-pigmented, protective membrane 

covering the gill arches and rakers. 

Haplochromis flaviijosephi 

Haplochromis flaviijosephi is the only non-African haplochromine, and has apparently to date 

never been seen in the aquarium hobby. That, however, is only because there have been no 

reports on the species in the relevant journals and magazines. It is in fact already being kept 

in the aquarium in Europe. I myself have been able to obtain it via two different Internet 

contacts, with the fishes originating from two sites close together in the vicinity of the Nahal 

Ha’Kibbutzim in Israel. 

The maximum size of males of this species is purportedly almost 13 centimetres (TL), but as a 

rule they attain only some seven centimetres in length. One of the specimens I studied had only 

26 scales in the lateral line, fewer than GREENWOOD (1979) cites as normal for Astatotilapia. 

GREENWOOD (1979) gives a size criterion of 70 mm SL for the change from bicuspid to unicuspid 

teeth in the outer series of the jaws. Because H. flaviijosephi only exceptionally attains a larger 

size, and I have never yet seen really large specimens of the species, I cannot provide any data 

on the tooth form of larger specimens. The specimens I studied all possessed bicuspid teeth, 

in which the anterior teeth in each case had the median cusp greatly protracted and the minor 

cusp was barely visible without pushing back the flesh of the gum in which it was partially 

embedded (swollen as a result of preservation?). The more lateral the teeth, the more they 

assume a subequilateral form. Among them was one tooth whose second cusp was so 

insignificant that the tooth gave the impression of being unicuspid. It may thus be that the 

minor cusp disappears (through wear?). The pharyngeal teeth are a real arsenal of different 

tools. In the centre, in the upper series, there are submolariform teeth, which become ever 

more delicate in form the closer they become to the edges of the bone. The flank scales are 

typical as given for Astatotilapia (see LIPPITSCH) (and in this respect, as already mentioned, she 

differs widely from GREENWOOD), ie there are very small ctenii and only on the outermost edge 

of the exposed part of the scale. The central ring structure of the scale is barely visible, as it 

is obscured by flattened granules and tubercles. The scalation on the breast is likewise typical, 

with only a gradual transition in size from the flank scales. 

The gill rakers are, unlike in H. desfontainesi, relatively uniform and triangular. 


21

Haplochromis sp. “Ismailia” 

In the aquarium males of this species attain a length of barely eight centimetres (SL). On 

looking into the mouth the distinctive discoloration of the tooth pulp is immediately apparent 

through the highly transparent enamel. Adult males and females can again be distinguished 

by the different form of the tooth crowns in the outer series. Those of males are 

predominantly unicuspid, those of females bicuspid, and in the latter there is sometimes a 

flange visible on the longer cusp. The bicuspid teeth are more close-packed than the 

unicuspid. 

There are two possible reasons for this sex-linked tooth form. Firstly, males and females may 

have different feeding habits. There is clear evidence for this in a work by SPATARU & GOPHEN 

(1985) on Haplochromis flavii josephi, according to which males with a length of 70 - 80 mm 

upwards feed largely on molluscs (probably snails by preference). In young males these 

constitute only 20-30% of gut contents. According to the same study, females feed exclusively 

on insect larvae (chironomids), amphipods, and oligochete worms. It would be interesting to 

Jaw teeth of a male (above) and female (below) H. sp. “Ismailia” (in each case the anterior right-hand side). Striking features include not 

only the variable form of the tooth crowns but also the different intervals between the individual teeth and the dark coloration of the 

tooth pulp, possibly mineral in origin and not previously known from any other species. 

22 


investigate whether the change in the jaw 

dentition requires the relevant food as a 

trigger, as has been established in the case of 

the pharyngeal dentition in Astatoreochromis 

alluaudi (HOOGERHOUD 1986). The other 

possible reason is at present no more than a 

hypothesis. In an earlier work (SCHRAML & 

TICHY 2010) I have surmised that males with 

unicuspid teeth are at an advantage during 

territorial battles, because they are better 

armed. But this remains a matter for more 

detailed investigation. 

The flank scales in H. sp. “Ismailia” possess 

more series of ctenii on the free margin of 

each scale than in the preceding two species. 

Right: Flank scale of Haplochromis sp. “Ismailia”.. 

Below: Magnified details of flank scale showing: (left) the ctenii, 

which in H. sp. “Ismailia” are arranged around the entire free 

margin of the scale, and in more series and closer together than in 

H. desfontainesi and H. flaviijosephi; and (right) the part normally 

covered by an adjacent scale, showing the radii. The very short 

tongue terminating the interradial zones is a striking feature. 


23

The form of the granulation is again irregular and blunt and largely obscures the central ring 

structure, although it is to be found on only around a third of the scale as a whole. The 

tongues at the margins of the interradial areas are narrower, but on the other hand the radii 

extend to the centre of each scale. The number of scales in the lateral line is, at 29, within 

the expected range of less than 30 for 

fluviatile haplo chromines (the fact that 

this is more than the 27 scales in the 

median series relates to the relatively 

large body depth). The head in this 

species is significantly broader than in 

H. flaviijosephi, and males possess far 

fewer eggspots (which are also often 

relatively large) than in the other two 

species. All three species have the 

common feature that, depending on 

mood, they may exhibit the same 

Female H. sp. “Ismailia”: mood-related pattern where horizontal stripes 

predominate (vertical striping weakly visible). 

melanin pattern, consisting of a median 

and a subdorsal stripe. 

It is anticipated that a more precise description of characters will follow when other 

populations/species found in Egypt are compared. 

References: 

BOULENGER, G. A. (1899): A revision of the African and Syrian fishes of the family Cichlidae. Part II. Proceedings of the Zoological Society of 

London, 1899 (1): 98-143, Pls. 11-12. 

DAGET, J. , GOSSE, J.-P., TEUGELS, G. G. & THYS VAN DEN AUDENAERDE, D. F. E. (1991): Check-list of the freshwater fishes of Africa. CLOFFA. 

ISBN Bruxelles, MRAC Tervuren, ORSTOM Paris. 4: i-xii + 1-740. 

GENNER, M. J. & HAESLER, M. P. (2010): Pliocene isolation of a north-west Saharan cichlid fish. Journal of Fish Biology, 76 (2): 435-441. 

GOPHEN, M. & SPATARU, P. (1985): Food composition and feeding habits of Astatotilapia flaviijosephi (LORTET) in Lake Kinneret (Israel). 

Journal of Fish Biology, 26 (5): 503-507. 

GREENWOOD, P. H. (1979): Towards a phyletic classification of the `genus' Haplochromis (Pisces, Cichlidae) and related taxa. Part I. Bulletin of 

the British Museum (Natural History) Zoology, 35 (4): 265-322. 

GREENWOOD, P. H. (1979): Towards a phyletic classification of the `genus' Haplochromis (Pisces, Cichlidae) and related taxa. Part II; the species 

from lakes Victoria, Nabugabo, Edward, George and Kivu. Bulletin of the British Museum (Natural History) Zoology, 39 (1): 1-101. 

GREENWOOD, P. H. & GEE, J. M. (1969): A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae). Part VII. Bulletin of the 

British Museum (Natural History) Zoology, 18 (1): 1-65. 

HOOGERHOUD, R. J. C. (1986): Ecological Morphology of some Cichlid Fishes. 133 p. Thesis, Rijksuniversiteit Leiden. 

ICZN (1999): International Code of Zoological Nomenclature, 4th edition. The International Trust for Zoological Nomenclature, London. i-xxix 

+ 306 pp. 

LACEPÈDE, B. G. E. (1802): Histoire naturelle des poissons. Plassan, Paris, France. 4: i-xliv + 1-728. 

LIPPITSCH, E. (1990): Scale morphology and squamation patterns in cichlids (Teleostei, Perciformes): A comparative study. Journal of Fish 

Biology, 37 (2): 265-291. 

LIPPITSCH, E. (1993): A phyletic study on haplochromine fishes (Perciformes, Cichlidae) of East Africa, based on scale and squamation 

characters. Journal of Fish Biology, 42: 903-946. 

MEYER, A. (1993): Phylogenetic relationships and evolutionary process in East African cichlids. Trends in Ecology and Evolution, 8: 279-284. 

MEYER, A., KOCHER, T. D. & WILSON, A. C. (1991): African fishes. Nature, 350: 467-468. 

PELLEGRIN, J. (1904): Contribution à l'étude anatomique, biologique et taxonomique des poissons de la famille des Cichlidés. Memoires Societe 

Zoologique de France, 16 (2-4): 41-400. 

REGAN, C. T. (1922): The classification of the fishes of the family Cichlidae. II. On African and Syrian genera not restricted to the great lakes. 

Annals and Magazine of Natural History, (9) 10 (57): 249-264. 

SCHRAML, E. (2007a): Haplochromis desfontainii. Blaulippenmaulbrüter. Die Aquarien- und Terrarienzeitschrift (DATZ), 60 (4): 62-66. 

SCHRAML, E. (2007b): Der Blaulippenmaulbrüter Haplochromis desfontainii. Historie und Haltung einer vom Aussterben bedrohten Art. DCG- 

Informationen, 38 (11): 241-250. 

SCHRAML, E. & TICHY, H. (2010): A new species of Haplochromis, Haplochromis katonga n. sp. (Perciformes: Cichlidae) from the Katonga 

River, Uganda. aqua, International Journal of Ichthyology, 16 (3): 81-92. 

SNOEKS, J. (1994): The haplochromines (Teleostei, Cichlidae) of Lake Kivu (East Africa): a taxonomic revision with notes on their ecology. 

Annales du Musée Royal de l'Afrique Centrale (Zoologie), 270: 1-221. 

SPATARU, P. & GOPHEN, M. (1985): Food composition and feeding habits of Astatotilapia flaviijosephi (LORTET) in Lake Kinneret (Israel). 

Journal of Fish Biology, 26: 503-507. 

VAN OIJEN, M. J. P. (1996): The generic classification of the haplochromine cichlids of Lake Victoria, East Africa. Zoologische Verhandelingen 

(Leiden), No. 302: 57-110. 

WITTE, F. & WITTE-MAAS, E. L. M. (1987): Implications for taxonomy and functional morphology of intraspecific variation in Haplochromine 

cichlids of Lake Victoria with descriptions of five zooplanktivorous species. Pp. 1-83, Tables 1-25. In: F. Witte, From form to fishery: an 

ecological and taxonomical contribution to morphology and fishery of Lake Victoria cichlids. Ph.D. Thesis Rijksuniversiteit, Leiden. 

24 


One of the German terms for cichlids is Buntbarsche, one of the few German words also 

known in the English-speaking world, at least among enthusiasts. The American Cichlid 

Association for instance, has named their journal Buntbarsche Bulletin. 

Among many German enthusiasts there is not really any clear distinction between the terms 

Barsch (perch) and Buntbarsch (colourful perch) and when a club member speaks of “meine 

Barsche” (my perches) he doesn’t as a rule mean that he is actually keeping members of the 

family Percidae (perches) such as the European Perch (Perca fluviatilis), for example, but is 

in all probability referring to cichlids (family Cichlidae). But despite certain external 

similarities between the two (one is minded in particular of the South American genus 

Cichla), the two families aren’t in fact so dreadfully close at all in phylogenetic terms. 

The Percidae (perches) belong to the suborder Percoidei (perches and their relatives) of the 

order Perciformes (perch-like fishes) in the superorder Acanthopterygii (spiny-finned fishes). 

Until recently the cichlids were included in the suborder Labroidei (wrasses and their allies), 

which, of course, again belongs to the order Perciformes (perch-like fishes). But according to 

a molecular genetic study published last year by LI and his colleagues, in which a new nuclear 

DNA marker (RNF213) was utilised, the phylogenetic history of the cichlids has to be rewritten 

and they belong to neither the Labroidei nor the Perciformes. According to this study 

the cichlids belong to a separate phylogenetic branch, together with the Atherinomorpha 

(smelts and their allies), which also includes the Cyprinodontiformes (toothcarps), for 

example the livebearing (Poeciliidae) and egg-laying toothcarps, the Mugiloidei (mullets and 

their allies), the Plesiopidae (spiny basslets), the Gobiesocoidei (clingfishes and their allies), 


Cichlids (and other acanthopterygian fishes) 

phylogenetically reclassified 

by KURT F. DREIMÄTZ 

LI et al. used two Haplochromis species for their genetic research. One of them was supposedly H. nubilus. 

25

and the Pomacentridae (damselfishes). Hitherto just a single morphological characteristic 

has linked all these groups of fishes, namely their eggs. The special characteristic of these 

eggs is sticky threads (“adhesive chorionic filaments”) around the opening (micropyle) by 

which sperm enters. 

LI et al. remove both the cichlids and the Pomacentridae from the Labroidei (wrasses and 

their allies), and, instead of erecting a new suborder for the new group, create a new order 

which they term the Stiassnyformes. Eggs such as are found in the above-mentioned groups 

are also known from other groups of fishes that LI et al. have not investigated but which they 

now consider to be possible members of their Stiassnyformes. These are the Pseudo - 

chromidae (dottybacks), Opisthognathidae (jawfishes), and Grammatidae (basslets). These 

and the Pholidichthyidae (convict blennies) had previously turned up as a phylogenetic 

lineage in an earlier study by SMITH & CRAIG (2007), but the latter authors did not cover all 

the spiny-finned fishes dealt with in the current study. However, the Gobioidei (gobies and 

their allies) and the Kurtidae (nurseryfishes) also possess such eggs, but according to the 

study by LI et al. belong to another phylogenetic lineage. On the other hand, the 

Embiotocidae (surf perches) and Mugiloidei (mullets and their allies) do not have such eggs 

but have been found to be members of the Stiassnyformes on the basis of DNA markers. The 

authors do not offer any credible explanation for these anomalies. A further anomaly in their 

work is that it creates another new order, the Blenniiformes (blenny-like fishes), which is 

supposedly part of the Stiassnyformes. An order part of another order??? 

Overall the Acanthopterygii (spiny-finned fishes) have been subjected to a huge reorganisation. 

Additional new taxa created by LI et al. at higher levels include the Zeioi - 

gadiformes, Stromateoidei, Anabantiformes, Paratrachinoidei, Zoarciformes, Cottimorpha, 

Triglimorpha, Serraniformes, Carangimorpha, and Epigonoidei. It remains to be seen 

whether the results of the research are confirmed by other scientists or whether there are 

other possible interpretations. Specifically, obvious differences in phenotype, for example 

between toothcarps and cichlids, are in no way explained by the new grouping. 

The International Code for Zoological Nomenclature does not regulate strictly the treatment 

of taxa above superfamily and lays down only very sketchy precepts, for example that the 

name must be uninominal (consisting of a single name). There is, however, no precept 

requiring that a subordinate taxon should be designated as the type of a superior taxon above 

the level of superfamily. There is also no requirement to indicate the intention to create a new 

name by an explicit additional phrase, as is expressly prescribed for a new species (n. sp.), 

new genus (n. gen.), and other taxa up to the level of superfamily. It would nevertheless be 

sensible to do so – but LI et al. have not. 

As a rule it is customary at the higher levels of taxonomy to name a superior category after 

a well-known representative, which in this case might have meant the Cichlidiformes, for 

example. Not so in this case. LI et al. have named an entire order after a living ichthyologist, 

MELANIE L. J. STIASSNY, the Curator of the Department of Ichthyology at the American Museum 

of Natural History. What an honour! 

References: 

ICZN (1999): International Code of Zoological Nomenclature. Fourth edn. The International Trust for Zoological Nomenclature, London. i-xxix + 

306 pp. 

LI, B., DETTAÏ, A., CRUAUD, C., COULOUX, A., DESOUTTER-MENIGER, M. & LECOINTRE, G. (2009): RNF213, a new nuclear marker for 

acanthomorph phylogeny. Molecular Phylogenetics and Evolution, 50: 345–363. 

SMITH, L. & CRAIG, M. (2007): Casting the percomorph net widely: the importance of broad taxonomic sampling in the search for the placement 

of serranid and percid fishes. Copeia, 2007 (1): 35–55. 

26 DREIMÄTZ: Cichlids reclassified

Katonga is the name of a river in Uganda and more recently also that of a Haplochromis 

species to date found only in this water. The Katonga doesn’t really deserve the name of river 

at all, as it is largely choked with papyrus and reeds. The swamp in which the Katonga rises, 

and which it drains into Lake Victoria to the east, is also drained by the Mpanga westwards 

to Lake George. The Katonga and Mpanga thus theoretically constitute a connection between 

Lake Victoria and lakes George and Edward. However, GREENWOOD (1973), wrote that the 

upper course of the Katonga was uninhabitable for all fishes except those that breathe air - 

and thus implied that this river did not permit any genetic exchange between Lake Victoria 

and lakes George and Edward. 

During an expedition in 1998 HERBERT TICHY and ERWIN SCHRAML fished in the Katonga not far 

from its source and despite GREENWOOD’s statement caught gill-breathing fishes there – small 

barbs (Barbus sp.), Pseudocrenilabrus victoriae, Oreochromis sp., and a Haplochromis 

species that could not be assigned to any previously described taxon. Because it was the only 

The holotype of H. katonga, the only adult male of the species found to date, shortly after capture. 

Haplochromis known from this river to date, they described the species as new, choosing the 

name of the river as the specific name and Haplochromis as the genus name. In so doing they 

followed the advice of VAN OIJEN, who has recommended choosing this generic designation for 

all haplochromine cichlids from the Lake Victoria basin for the time being. In fact the new 

species cannot be unequivocally assigned to any of the genera erected or revalidated by 

GREENWOOD, as it exhibits affinities with Astatotilapia as well as with Enterochromis. 


A new Haplochromis described from the Katonga 

by FRANK SCHÄFER 

27

The fishes were relatively uncommon at the collecting site near the hamlet of Kabagole. 

Despite an extensive search, during which large parts of a channel cleared for canoes were 

fished with a drag net, only nine specimens came into the hands of the describers, and only 

one of those an adult male. 

Genetic studies have shown that this species is more closely related to species from Lake 

Victoria than to those from lakes Edward and George or other haplochromines. At less than 

seven centimetres (SL), the species remains relatively small. Because male and female 

cannot unequivocally be assigned to a single trophic group and no studies were undertaken 

One of the females of H. katonga from the type series. 

of the food consumed, it can only be surmised that these fishes are either an insect-eating 

species or that they feed on zooplankton. The heavy black deposits on the jaw teeth 

indicate a particular, as yet unclarified dietary preference. The bold red coloration on the 

anal and caudal fins, the iridescent greenish colour of the flanks, and the bluish-grey head 

and dorsum in males are very distinctive and would also make it an attractive aquarium 

fish. A pity that the species has to date not been imported alive. 

References: 

GREENWOOD, P. H. (1973): A revision of the Haplochromines and related species (Pisces: Cichlidae) from Lake George, Uganda. Bulletin of the 

British Museum of Natural History (Zoology), 25 (5): 141-242. 

SCHRAML, E. & TICHY, H. (2010): A new species of Haplochromis, Haplochromis katonga n. sp. (Perciformes: Cichlidae) from the Katonga 

River, Uganda. aqua, International Journal of Ichthyology, 16 (3): 81-92. 

VAN OIJEN, M. J. P. (1996): The generic classification of the haplochromine cichlids of Lake Victoria, East Africa. Zoologische Verhandelingen 

(Leiden), No. 302: 57-110. 

28 

SCHÄFER: New Haplochromis described

Anyone who has had anything to do with the subject of taxonomy will be aware that the 

grammatical gender of a genus name can have an effect on the species name, specifically 

when the latter takes the form of an adjective. Grammatical gender is a feature of most 

languages, especially those of Europe, and is even found in Old English (Anglo-Saxon), 

albeit not in modern English. 

The German reader of 

eggspots will not, however, 

derive any great benefit from 

this in terms of under - 

standing, as the Latinderived 

genders involved 

here are significantly 

different to the German. 

As always where taxonomy is 

concerned, the Code of the 

International Commission on 

Zoological Nomenclature 

(ICZN) provides relevant 

precepts, and in this case in 

particular, helpful examples 

as well. Although these rules 

occupy several pages in the 

Code (and hence repeating 

them in full in this article 

would take too much space), 

the examples nevertheless 

cannot hope to explain 

everything relevant, and as a 

result authors without any 

special knowledge of Latin 

often make mistakes. The 

commonest is that they do 

not appreciate when a 

species name takes the form 

of a noun and when it 

consists of an adjective. 

Examples of this include 

names that end in –taenia, 

which means ‘band’ and 

hence is a substantive, while 

the ending taeniatus (-ata, - 

atum) signifies ‘banded’ and is, of course, an adjective. In fact the matter can be even more 

complicated, as names that end in, for example, -urus can be either a substantive or an 


The gender of the genus Haplochromis 

and other –chromis 

by MARY BAILEY and ERWIN SCHRAML 

Chromis chromis (UW-Photo Adriatic Sea), the species from whose name the suffix 

commonly used for cichlid genera is derived. 

29

adjective at the discretion of the author of the name, and it is common for such names to 

be incorrectly inflected as adjectives when they are actually substantives. For this reason 

the ICZN also advises that every new description should not only give an indication of 

etymology but also state the gender of a name in the case of a new genus, and what type 

of word the name is in the case of a new species. 

You may well ask what all this has to do with cichlids whose genus names end in –chromis. 

Well, if you browse through ESCHMEYER’s Catalog of Fishes for genera with such an ending, 

you will find that they are all designated as masculine. It is also laid down by the Code that 

the last part of a compound name determines its gender, and because all these compound 

genus names are formed by the simple addition of -chromis to the first part of the name, it 

is this suffix that determines the gender. 

It is also important to realise that this suffix, in the taxonomic sense, is derived not from 

the Ancient Greek word chroma meaning ‘colour’, but from a marine fish with this name in 

Ancient Greek. And in fact for a long time various cichlids were assigned to the genus 

Chromis (if you search for the terms Chromis and Cichlidae in the Catalog of Fishes you 

will get a list of 84 relevant entries!), which still exists today but is now reserved for reef 

fishes of the family Pomacentridae (formerly known as the Chromidae). This is the reason 

why, for example, Haplochromis was originally erected as a subgenus of Chromis (for the 

species Chromis (Haplochromis) obliquidens). 

The final part of the equation is that in 1986 the genus Chromis was designated as feminine 

by an official act of the ICZN (Opinion 1417). This means that in consequence a number of 

familiar names, for example Chromis chrysurus and C. flavomaculatus, are now C. chrysura 

and C. flavomaculata respectively. 

From this one might conclude that in consequence all genus names that relate to Chromis 

and include this taxonomic suffix would likewise have to be changed, and assume that 

many African cichlid genera as well as various genera of marine fishes would also be 

affected. But don’t worry! The ICZN, whose main objective is the stability of nomenclature, 

has put the brakes on the avalanche of name changes that would otherwise inevitably have 

followed. It is expressly stated in Opinion 1417 that “It is to be noted that this in no way 

defines the gender of generic names ending in –chromis”. Phew! So we don’t have to 

relearn everything after all! 

References: 

ESCHMEYER, W. N. (ed.) Catalog of Fishes electronic version (19 February 2010). 

http://research.calacademy.org/ichthyology/catalog/fishcatmain.asp 

ICZN (1986): Opinion 1417. Chromis CUVIER in DESMAREST, 1814 (Osteichthyes: Perciformes); gender confirmed as feminine. Bulletin of 

Zoological Nomenclature, 43 (3): 267-268. 

ICZN (1999): International Code of Zoological Nomenclature. Fourth edn. The International Trust for Zoological Nomenclature, London. i-xxix + 

306 pp. 

30 BAILEY & SCHRAML: Gender of -chromis

Dissolved through lack of interest! These few words, brief and to the point, and dire 

to the enthusiast’s ear, mark the end of an era. Many aquarium clubs are battling 

against the constant loss of members. But even so it is a shock to hear that a national 

organisation has found itself obliged to fold. The more so in that the organisation in 

question is one whose dynamism has brought it fame beyond the borders of its own 

country. Yet until just a few years ago it regularly organised a major aquarium show 

in Antwerp. For many clubs and regional groups of other cichlid organisations the 

“Cichlidenshow” was a reason to visit the city, sometimes by the busload. The 

members of the Belgische Cichlidenliefhebber Vereniging (A. B. C. V. = Belgian 

Cichlid Association) put on this incomparable event, sometimes every other year, in a 

former cinema in the 

suburb of Borgerhout. The 

aquaria, some of them 

constructed of wood, were 

always attractively and 

often also innovatively 

decorated. The fishes, 

cichlids from all over the 

world, were impressive, 

and frequently spectacular 

and rare. Many an 

illustrious guest attended 

the exhibition, and note - 

worthy experts were 

always invited to be guests 

at the assemblies of the 

association. Cichlidae, the 

informative journal of the 

association, was indepen - 

dently published even in 

the early years. In its 

heyday the club had 

several hundred members, 

but only five turned up at 

the last member assembly, 

even though it was known 

that matters had reached a 

“life or death” situation. 

Given that sort of lack of 

interest, the “last of the 

Mohicans” had no choice 

but to dissolve the 

association on 22nd 

October 2009, after more 

than 35 years in existence. 


The end of an era 


View from the upper floor of the former cinema, showing some of the aquaria at the (9th) 

“Cichlidenshow” for which the Belgian Cichlid Association was famous far and wide. 

31

A little known Pseudocrenilabrus species from East Africa 

by RICO MORGENSTERN 

It was a good ten years ago that I first saw an unfamiliar Pseudocrenilabrus species 

on sale in a Chemnitz pet shop as Pseudocrenilabrus philander. Interestingly there 

was just one male among the 20 or so individuals, and I bought it along with three 

females. These fishes exhibited certain similarities to both P. philander and P. 

victoriae, but I came to the conclusion that they must be another species. And initially 

I was unable to discover any more about them, so until further information became 

available all I could do was hope that they would grow well and breed. Unfortunately, 

however, this was not the case: two females died after only a few days, the remaining 

pair spawned several times and the eggs developed, but I was unable to rear any 

youngsters, as they proved very delicate. And finally an outbreak of the normally easyto-treat 

Ichthyophthirius (or the treatment used?) led to the loss of the fishes. 

Enquiries at the shop where I had purchased the fishes failed to provide either 

information on their origins or the opportunity of obtaining more. I first found photos 

of the species in LAMBOJ (2004), where on page 221 a male and female of unknown 

provenance are depicted as Pseudocrenilabrus philander. A short time later I came 

across an interesting article by SEEGERS (1996a), which not only pictured “my” fish but 

also gave its provenance: apparently it originated from the upper Ruaha drainage in 

Tanzania (see below). 

Pseudocrenilabrus sp. "Ruaha", male in courtship coloration. 

32 

MORGENSTERN: Pseudocrenilabrus sp. "Ruaha"

In February 2006 I finally again saw live specimens. Once again the females were in the 

majority, but at least there were several males this time. Nevertheless I bought just one 

pair, as I had learned in the course of time that in the case of Pseudocrenilabrus species 

and some other haplochromines it is unwise to keep a single male with a “harem” because 

of the aggression of the females among themselves. Because the weakest individual is 

then attacked by all conspecifics, the desired effect of spreading the aggression often 

becomes precisely the opposite. 

Male Pseudocrenilabrus sp. "Ruaha" enticing the female to the spawning site. 

By and large these fishes can be maintained like other Pseudocrenilabrus, but my initial 

fear that the species was somewhat more sensitive and delicate was to be confirmed. 

Initially, however, the fishes did quite well and spawned regularly, but it took four 

attempts before the female finally brooded to term. Unfortunately the fry took hardly any 

food and died fairly quickly. At the next attempt only bellysliders were produced - the 

reasons for this are unknown, but I had previously also had this problem with P. 

multicolor. Perhaps it related to the condition of the female, so I separated the fishes for 

a number of weeks. 

When the female’s genital papilla was clearly visible I put the male back with her, and he 

immediately began to court her. I was now able to observe the courtship and spawning 

behaviour in full detail. The male approached the female in an unusual way. He adopted 

a diagonally downward-pointing position, with all his fins erect. Apart from the rapidly 

fanning pectoral fins the fish looked as if he had been struck rigid as he slowly moved 

towards the female. When he reached her he began typical quivering behaviour, 

displaying in front of the female in a slightly curved position, dorsal fin slightly folded, 

anal fin spread, but not extended towards the female in the way with which I was familiar 


33

from P. victoriae and P. philander and which can be seen even more strikingly in species 

with eggspots. The final stage was typical leading behaviour by the male, to guide the 

female in the direction of a shallow pit he had excavated previously, and which he 

repeatedly improved. The entire sequence was repeated numerous times until the female 

followed to the “nest”. The spawning process resembled that of the other 

Pseudocrenilabrus, and will be described and discussed in more detail elsewhere. 

After spawning I placed the female in a small rearing tank where she could incubate her 

brood undisturbed. After 12 days (at a constant temperature of 26 °C) the young were 

released for the first time, and this time were normally developed. There were around 40 

of them. The fry were taken back into the mouth for another four days at night and when 

danger threatened, but the female then ceased brood care so I caught her out. The fry fed 

well right from the start, but took only Artemia nauplii and sieved pond foods; flake food 

was accepted only after a week. They grew well, and hence I hoped that this time I would 

succeed in rearing them. After six weeks they already measured a good two centimetres 

long and were already exhibiting some of the typical markings of the species. But then I 

spotted a number of individuals with noticeable respiratory distress, and shortly thereafter 

there were occasional corpses lying in the aquarium, then increasingly more, and after a 

week the whole brood was dead. I was filled with despair. The cause remains a puzzle, the 

water parameters were as they should be. And to cap it all the female had now also become 

unwell, she was no longer able to swim in the open water but merely “crawl” around on the 

bottom. Another breeding attempt was obviously out of the question. 

I subsequently managed to obtain individuals of the species on two further occasions (most 

recently in summer 2008), but in each case they were already rather sickly at the time of 

purchase and died soon afterward, without producing any offspring. And that was the end 

of that chapter, but I think that there may still be aquarium strains in existence, as these 

fishes keep turning up and are also sometimes available simultaneously in a number of 

different shops. 

Provenance 

According to SEEGERS (1996a), this fish originates from the Usangu Plain in the upper 

Ruaha drainage in Tanzania. More precise details of the location can be found in SEEGERS 

(1996b: 716) (translated from German): “Finally, it turned out that the Kimani River is 

crossed by the road to Dar es Salaam around 95 kilometres east of Mbeya; however, there 

are no waterfalls there. But there may well be a fast-flowing stretch of river [...]. Nowadays 

a weir has been constructed there and part of the water diverted for irrigating rice crops. 

Trial collections demonstrated that not only does Parakneria tansaniae occur regularly at 

this spot, but also that numerous other fishes are to be caught there, such as various barbs, 

Labeo cylindricus, Mesobola spinifer, Amphilius jacksonii, Leptoglanis rotundiceps, 

Chiloglanis deckenii, Haplochromis sp., and Pseudocrenilabrus sp. A very large number of 

the species found here are rheophilic [...], even though the current is certainly not very 

turbulent (in the most literal sense of the word)”. 

SEEGERS (pers. comm.) found the species only at this site, and I have so far been unable to 

find any other evidence regarding this or additional locations. There are no details known 

of how these fishes came to be in the trade. They might be linked with the specimens 

brought back by SEEGERS, but it is also possible that they have been imported several times, 

as killifish enthu siasts have now found their way into the area. For unknown reasons, 

34 MORGENSTERN: Pseudocrenilabrus sp. "Ruaha"

enqui ries as to the 

provenance of these fishes 

in various shops have not 

been answered with any 

great willingness, though in 

one case I did manage to 

elicit the information that 

they came from a Czech 

breeder or wholesaler. 

Even though the precise 

origins of the fishes I kept 

thus remains unclear, I 

nevertheless believe that 

their very distinctive 

characteristics leave no 

doubt that they were the 

same form. I am of the 

opinion that they do not 

belong to P. philander in the 

broad sense. I am far more 

inclined to regard them as a 

well-defined and distinct 

species, for which I suggest 

using the provisional name 

of Pseudocrenilabrus sp. 

“Ruaha”. 

Characteristics 

This is a rather slender, elongate fish, which in my experience attains a total length of 

around eight centimetres; females re main somewhat smaller. The upper head profile is 

slightly rounded, the mouth relatively small. The dorsal fin is rather low, and the soft-rayed 

part is pointed, at least in males. The first ventral-fin ray is no or only slightly longer than 


Above: Spawning procedure is as in 

other Pseudocrenilabrus, here the Tposition. 

Left: Female with larvae in her mouth. 

Below: For several days after first release 

of the brood, the fry are taken back into 

the female’s mouth when danger 

threatens. 

35

the second, so that the form of ventral fins often appears almost rounded, especially in 

females. 

The markings are very characteristic; they consist of a longitudinal band, around two scalerows 

wide anteriorly, tapering to one scale-row wide on the caudal peduncle. The posterior 

part (from around the level of the spinous part of the anal fin) is continuous, while on the 

anterior body there are regular lighter areas, creating the impression of a row of spots. 

Depending on mood, these spots can in turn become part of a pattern of crossbands, most 

clearly seen in females during brood care. Dominant males almost always exhibit a dark 

(lachrymal) stripe on the snout. 

Males have a brownish base coloration, paling to yellowish on the flanks and white on the 

underside. The scales on the upper half of the flank and on the caudal peduncle are metallic 

bluish green, those on the mid-flank golden, and those on the belly light blue. The lower lip 

is bright blue, a narrow stripe of the same colour extends along the lower margin of the 

cheek to the preoperculum, and there are a number of additional irregular metallic blue 

markings on the lower part of the operculum. The dorsal fin has a blackish margin, 

somewhat broader anteriorly and tapering posteriorly, followed by a narrow red band and 

then a striking metallic blue stripe extending right to the tip of the fin. There are hints of 

this in many members of the genus, but I know of no other form in which it is so clearly 

and evenly expressed. The proximal part of the dorsal fin exhibits a double row of deep red 

spots on the spinous portion, which is covered in fainter blue markings; in the soft-rayed 

part there are alternating metallic blue and reddish stripes or rows of dots running 

diagonally across the fin-rays on a background of transparent yellowish green, becoming 

colourless distally. The upper part of the caudal fin is similar, but much more faintly 

coloured, in particular the blue dots are barely visible. The lower half of the caudal fin is 

yellowish, at its base there are several large, irregular, metallic blue dots, and with 

increasing age vertical rows of dots extend posteriorly. The anal fin exhibits broad metallic 

blue stripes separated by only narrow yellowish interspaces, a grey anterior margin, and a 

small, irregular red spot at the tip. The ventral fins are reddish in colour apart from the 

anterior margin. 

During courtship the coloration intensifies, the sides of the head and body are now bold 

golden yellow, a number of scales above and behind the pectoral fins have red centres, and 

the red on the fins, especially the ventral fins, becomes more intense. Interestingly the 

sooty black coloration seen on the throat and breast, as well as the on the ventral fins, in 

many other species is almost completely absent in this case. 

The females resemble those of P. victoriae and some forms of P. philander in their 

coloration, but are can easily be distinguished on the basis of their typical markings. Traces 

of the striking dorsal-fin markings in males are also apparent in females, and the intensity 

of the blue head pattern that brightens up females during brood care also appears to be 

diagnostic. 

Comparison with other species 

So how is this species to be classified taxonomically? To date two members of the 

Haplochromis assemblage have been described from the Rufiji-/Ruaha system, namely 

Paratilapia kilossana STEINDACHNER, 1914 and P. vollmeringi STEINDACHNER, 1914. The types 

are adult specimens; one syntype of P. kilossana originates from Kilosa (Wami system), the 

36 MORGENSTERN: Pseudocrenilabrus sp. "Ruaha"

second, as well as the holotype of P. vollmeringi, from Kidatu on the Ruaha. In the light of 

current knowledge the detailed descriptions and illustrations in STEINDACHNER (1916) leave 

little doubt that they are male and female of the same species, although REGAN (1922) 

synonymised P. kilossana with Haplochromis bloyeti and P. vollmeringi with H. gigliolii. He 

regarded the latter species as “perhaps not distinct from H. moffati” (= Pseudocrenilabrus 

philander). However the description of the holotype of Hemichromis gigliolii PFEFFER, 1896 

from the Kingani River (nowadays the Rufu) corresponds to that of an Astatotilapia species 

widespread in the Tanzanian coastal basin, which according to SEEGERS (1996c) should be 

assigned to A. bloyeti (SAUVAGE, 1883). The type locality is “Kandoa” (also Kondoa; 

according to old maps this place was situated in the vicinity of what is now Kilosa, Wami 

drainage, and is not to be confused with the modern Kondoa). Ctenochromis strigigena 

PFEFFER, 1893, described from Mbusini and Matamondo in the lower Wami drainage, has 

several times been confused with Pseudocrenilabrus species, and is today regarded by 

some authors as likewise a synonym of A. bloyeti. 

From this we may assume that we are dealing with an undescribed species. As far as is 

known at present its distribution is isolated; the closest populations geographically are to 

be found in the Malagarasi (Pseudocrenilabrus victoriae SEEGERS, 1990; for the status of the 

species see MORGENSTERN 2010), in the Wembere (P. cf. victoriae), in the Lake Rukwa 

system (only the upper Saisi drainage; P. cf. philander), as well as in the Lake Malawi basin 

(P. philander (WEBER, 1897)). It is noteworthy that this is the only Pseudocrenilabrus 

species found to date in an eastward-flowing system north of the Zambesi, if we discount 

P. philander from the Lake Chilwa/Chiuta basin (Ruvuma system) and a probably 

introduced population of P. victoriae in the Athi River (Kenya). 

P. victoriae resembles P. sp. “Ruaha” as regards the metallic scale pattern, which extends 

practically all over the exposed part of each scale, above all in the ventral region and the 

anterior part of the flank, unlike in P. philander where it is limited to the outer margin of 

the scales. In addition, both forms exhibit only a small amount of black in the ventral fins. 

P. victoriae is, however, less slender and has noticeably pointed ventral fins in both sexes, 

and in adult males these may even be prolonged into fairly long threadlike extensions. The 

form of the mouth is similar in P. victoriae, but the mouth is somewhat smaller. In addition 

both lips are coloured bright blue, while otherwise there are no blue head markings 

present. There is at best only a hint of the blue band in the dorsal fin, but on the other hand 

the caudal fin is much more intensely and evenly spotted with blue. The anal fin is more 

striped rather than spotted, but the pattern is less dense in this case. Finally, the large, 

bright orange-red spot on the tip of the anal fin is particularly characteristic. 

I regard the form from the Wembere/Kitangiri drainage in northern central Tanzania as 

rather similar, at least on the basis of the photo published by SEEGERS (1990, 1996a). In 

particular the dorsal fin pattern is highly reminiscent of that of the fish portrayed here, but 

the blue submarginal band is not so clearly expressed. The fish in the photo exhibits a 

similar head and body form, but is not quite as slender. It resembles P. victoriae as regards 

the blue lips, the anal-fin pattern, and the long ventral fins. 

It is difficult to make generalisations regarding differentiating characteristics when it 

comes to the multitude of different forms currently grouped together as P. philander. In this 

case reference must be made to the typical characters of P. sp. “Ruaha”, which are 

unknown from any other form, at least in this combination. P. philander usually has a more 

robust body form and a larger mouth, the dorsal band is absent or only weakly expressed, 


37

the anal fin is spotted rather than striped, and the ventral fins in reproductively active 

males are as a rule at least partially black pigmented. In addition they are usually pointed 

and the first soft ray is prolonged into a short threadlike extension, but this doesn’t apply 

in, for example, the form described as Pseudocrenilabrus sp. aff. philander from the upper 

Saisi drainage (Rukwa system) by SEEGERS (1996c); similar fishes are also found in the 

upper Chambeshi region and in southern affluents of Lake Tanganyika). This fish is, 

however, quite different in coloration and corresponds to other forms of P. philander as 

regards overall habitus. 

The form of the ventral fins in the Ruaha fishes is, however, interesting for another reason, 

particularly in connection with the slender form. Specifically, rounded ventral fins, in 

which the second ray is somewhat longer than the first, and body form – in addition to 

scalation characters – were the reasons why GREENWOOD (1979, 1984) assigned 

Orthochromis machadoi (POLL, 1967) a rheophilic cichlid from the Cunene in Angola, to the 

genus Orthochromis, although in terms of colour characters, including the genus-typical 

anal-fin pattern, it corresponded closely to the genus Pseudocrenilabrus. In the original 

description POLL had already indicated a close relationship with P. philander, and according 

to him the morphological characteristics shared with the species then assigned to the 

genera Orthochromis and Rheohaplochromis were the result of convergent evolution. This 

appears thoroughly plausible, as many of these characters are also found in rheophilic 

cichlids from unrelated groups (including New World species). In addition, according to 

DNA studies (KOBLMÜLLER et al. 2008) O. machadoi should be assigned to the genus 

Pseudocrenilabrus. In terms of certain morphological characters (only in relation to the 

expression of the characters, a close relationship is unlikely by virtue of the widely 

separated distribution regions) Pseudocrenilabrus sp. “Ruaha” may occupy a position 

between O. machadoi and the rest of Pseudocrenilabrus, but this is something that requires 

more detailed study. 

References: 

GREENWOOD, P. H. (1979): Towards a phylogenetic revision of the ‘genus‘ Haplochromis (Pisces, Cichlidae) and related taxa. Part I. 

Bulletin of the British Museum (Natural History) Zoology, 35 (4): 265-322. 

GREENWOOD, P. H. (1984): The Haplochromine species (Teleostei, Cichlidae) of the Cunene and certain other Angolan rivers. Bulletin of 

the British Museum (Natural History) Zoology, 47 (4): 187-239. 

KOBLMÜLLER, S., SCHLIEWEN, U. K. , DUFTNER, N., SEFC, K. M., KATONGO, C. & STURMBAUER, C. (2008): Age and spread of the 

haplochromine cichlid fishes in Africa. Molecular Phylogenetics and Evolution, 49: 153-169. 

LAMBOJ, A. (2004): Die Cichliden des westlichen Afrikas. Birgit Schmettkamp Verlag, Bornheim 

MORGENSTERN, R. (2010): Pseudocrenilabrus victoriae Seegers, 1990. www.cichlidae.com 

PFEFFER, G. (1896): Die Fische Ostafrikas. In: K. Möbius: Deutsch-Ostafrika, Vol. 3.: Die Thierwelt Ost-Afrikas und der Nachbargebiete. 

Wirbelthiere. Verlag Dietrich Reimer (Ernst Vohsen), Berlin. 3: i-xviii + 1-72. 

POLL, M. (1967): Contribution à la faune ichthyologique de l'Angola. Publicações. Culturais da Companhia de Diamantes Angola, 75: 1- 

381, Pls. 1-20. 

REGAN, C. T. (1922): The classification of the fishes of the family Cichlidae. II. On African and Syrian genera not restricted to the great 

lakes. Annals and Magazine of Natural History, (9) 10 (57): 249-264. 

SEEGERS, L. (1990): Bemerkungen zur Gattung Pseudocrenilabrus. 2. Pseudocrenilabrus multicolor victoriae nov. subspec. Die Aquarien 

und Terrarien-Zeitschrift (DATZ), 43 (2): 99-103. 

SEEGERS, L. (1996a): Kleine Maulbrüter aus Afrika: Die Gattung Pseudocrenilabrus. Das Aquarium, 30 (9): 6-12. 

SEEGERS, L. (1996b): Die Kneriiden Ost- und Zentralafrikas. Die Gattung Parakneria. Die Aquarien und Terrarien-Zeitschrift (DATZ), 49 

(11): 714-718. 

SEEGERS, L. (1996c): The Fishes of the Lake Rukwa Drainage. Annales de la Société Royale Zoologique de Belgique, 278: 1-407. 

STEINDACHNER, F. (1915): Bericht über die ichthyologischen Aufsammlungen der Brüder Adolf und Albin Horn während einer im Sommer 

1913 ausgeführten Reise nach Deutsch-Ostafrika. Denkschriften der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen 

Akademie der Wissenschaften in Wien, 92: 59-86 [1-28], Pls. 1-5. 

38 

MORGENSTERN: Pseudocrenilabrus sp. "Ruaha"

In an earlier issue of eggspots (no. 2) editor ERWIN SCHRAML (2009) eloquently 

put forward some arguments in the dispute regarding Maylandia and 

Metriaclima, but I think it is quite clear that his views accord with the 

proponents of the former. So here is the view from the other side of the fence. 

To understand why Maylandia is a nomen nudum, and why this is important, it is 

necessary to consider the fundamental objective of taxonomy, which is, to put it 

as simply as possible, one name per taxon (genus, species, etc), so that anyone 

anywhere who encounters the scientific name of an animal will know precisely 

what animal is meant. The rules say (again put as simply as possible) that the 

earliest name used is the correct one (“Principle of Priority”), provided the name 

is valid, and the ICZN Code of Nomenclature lays down various further rules 

(termed “Articles”) that define what makes a name valid or not. 

As SCHRAML has stated, Article 13 is the critical rule in the case of Maylandia, but 

to date I have yet to find any pro-Maylandia author who has considered the 

fundamental purpose of that rule, or, indeed, actually understood its wording. 

Much of the difficulty seems to me to hinge on linguistic difficulties, as most of 

the pro-Maylandia protagonists do not speak English or French as their mother 

tongue (if at all), while the Code is laid down in both languages and MEYER & 

FOERSTER’s paper was published in the latter. A short word on my own credentials 

here: English is my mother tongue and I have a degree in it, as well as working 

professionally as an editor for some years, making sure material – some of it 

scientific – says what its (often non-English) authors intend. I have also worked 

for almost 20 years as a translator of ichthyological and other biological French 

and German into English, so MEYER & FOERSTER’s paper holds no linguistic 

mysteries for me. Among the Maylandia supporters many have little or no French, 

and only CONDÉ & GÉRY (1999) were native French speakers – and, perhaps 

significantly, they took refuge in a quite different argument to everyone else – 

but more of that later. 

Differentiate and differentiate from 

Article 13 requires that a name must be linked to a particular taxon – eg genus, 

subgenus, or species – by the provision of information, in words, that defines the 

taxon. The point of this is to allow other workers, perhaps with a pile of 

specimens to identify and classify, to determine whether any particular specimen 

belongs to that taxon. In order for this to be possible the description MUST 

provide an unequivocal list of characters that together define that taxon and no 

other. This is the meaning of the verb “differentiate” as used in Article 13 (in all 

three of the 2nd, 3rd, and 4th editions!). “Differentiate” used in this context 


Echo 

Maylandia or Metriaclima – the case for Metriaclima 

by MARY BAILEY 

39

means “define”, a specialised usage, and is not the same as “differentiate from” 

– the commonplace, everyday usage which “Maylandia-ites” cite so regularly with 

regard to MEYER & FOERSTER’s alleged (see below) differentiation of Maylandia 

from Pseudotropheus williamsi. Differentiating taxon X from taxon Y tells us 

only how to tell that X is not Y; it does not tell us how to distinguish X from 

anything else (in this case other closely-related (ie mbuna) genera and 

subgenera), or how to determine whether any particular animal belongs to taxon 

X, which is what Article 13 and “differentiate” require. 

A tacit cry for help? Maylandia or Metriaclima – after so many years a conclusive decision would seem called for. 

To give a trivial example. You could differentiate me from the editor of eggspots 

quite easily as I am a lot smaller and don’t have a beard, but that probably 

applies to most of the women and a lot of the men on this planet, and it does not 

tell you a thing about who I am, what makes me MB rather than any other woman 

you might come across. But “female English writer on cichlids” would enable a 

lot of fishkeepers to make a fair stab at who you were talking about. 

With me so far? 

Unfortunately very few of the people who support Maylandia, even those whose 

mother tongue is English, seem to appreciate this semantic nicety, but I am in no 

doubt whatsoever that the authors of the Code DID, and that they intended 

“differentiate” to mean “define”, ie differentiate from ALL other similar taxa, not 

just one. The introduction to the Code is quite specific about its use of precise, 

unambiguous language to convey specific meanings. Moreover any other sense 

40 

Help! 

Maylandia or Metriaclima 

– you decide. 

BAILEY: the case for Metriaclima

of “differentiate” would render the rule pointless as it would not fulfil its 

purpose, ie to ensure that a name is valid only if it does its job in allowing others 

to identify specimens as belonging or not belonging to the taxon in question. 

Maylandia or zebra complex? 

SCHRAML has drawn attention to another relevant problem of interpretation, this 

time of the wording of MEYER & FOERSTER rather than the Code. To avoid readers 

having to refer back to his article I will again quote the relevant text (translated 

by myself and, I believe, accurately). 

“Some other species of Pseudotropheus, s. lat., resemble Ps. greshakei and differ 

from Ps. williamsi by having less regular internal rows of teeth and, at least in 

adults, containing many unicuspid, small pharyngeal teeth which are very closepacked 

posteriorly, and the melanin pattern on the body forming vertical bars 

which are more or less visible. Such species include Ps. zebra BOULENGER, Ps. 

aurora BURGESS, Ps. lombardoi BURGESS and Ps. livingstonii BOULENGER. The last 

two species have, in females and non-territorial males, a well-defined pattern of 

six vertical bars on the body, five of which extend into the dorsal and which 

diminish to the point of disappearing on the lower part of the flanks. Ps. zebra 

and the many related forms have vertical bars, as in Ps. greshakei, but much 

bolder. We suggest that this zebra complex should be included in the subgenus 

Maylandia.” 

Maylandia supporters assert that the characters given here constitute a 

description of Maylandia; SCHRAML cannot understand why supporters of 

Metriaclima refute this. I can only assume that the problem is again linguistic, as 

it is quite clear and totally unambiguous that the characters listed relate not to 

Ps. greshakei but to “Some other species”. These “other species” are 

subsequently listed in part and finally described as a “zebra complex”. This 

group does not include Ps. greshakei, it merely resembles that species, and that 

means nothing as a resemblance does not necessarily imply a close phylogenetic 

relationship, it can be the result of, for example, similar environmental pressures 

producing a similar result. 

But, Ps. greshakei is the type species of Maylandia. As the characters listed do 

not belong to its type species, they cannot possibly constitute a description of 

Maylandia. 

Again, Maylandia supporters argue that what the text means is “…….resemble Ps. 

greshakei and like it differ from Ps. williamsi by having…….”, in an attempt to 

include Ps. greshakei in the list. But the text does not say that, it is describing 

two separate and unrelated features of the “other species”. And it is what you 

say, not what you mean to say, that is critical in science. Were scientific texts 

open to individual reader interpretation then the result would be chaos, so the 

assumption must always be that what is said is what is intended. 

Note that in any case this list of characters that Maylandia supporters point to as 

a “description” of the subgenus do not unequivocally define anything as required 

by Article 13; they mainly differentiate the zebra complex from Ps. williamsi, 


41

which we have already seen is not a valid method of establishing what a taxon is, 

only a very limited what it is not. 

Moreover the authors themselves are unsure which species should be included in 

the group they are “describing” and even go on to say that this depends on the 

weight assigned to one character. 

“Qu'il faille aussi inclure Ps. livingstonii et Ps. lombardoi, depend de la valeur 

que l'on accorde au patron melanique.” (Whether or not Ps. livingstonii and Ps. 

lombardoi should be included depends on the importance that one assigns to the 

melanin pattern.) 

Astute readers will note the paradox – Maylandia supporters assert that MEYER & 

FOERSTER have differentiated (defined) Maylandia, but the authors themselves 

state that their “definition” is not concrete, and what you include depends on 

how important you consider one of the “defining” characters. But it was their job 

to decide on the relevance of that character. 

Ah, but Maylandia supporters might argue that the list of characters has only to 

“purport to differentiate“ (= define) the taxon, that is, state what the authors 

believe is a valid definition at the time of writing. Well yes, but that is because 

new species and new information may come along later and require the taxon to 

be redefined (as has happened with Metriaclima, hence the 2006 redecription by 

KONINGS and STAUFFER broadening the scope of the genus in the light of additional 

knowledge). The Code cannot reasonably require that the description of a taxon 

does define it unequivocally, as a biologist can work only with the information 

currently available, which may not be complete. Hence biologists are merely 

required to provide what they believe to be a valid definition. But MEYER & 

FOERSTER’s “description” doesn’t do anything of the kind, as they themselves 

immediately question its validity on the basis of information that was available 

but which they hadn’t properly considered. 

To sum up, MEYER & FOERSTER actually provide a very sketchy description of the 

“zebra complex”, not Maylandia, and aren’t sure what species actually belong in 

that complex. They actually exclude Ps. greshakei (the type of Maylandia) from 

the zebra complex; and to cap it all, they aren’t even sure if this complex does 

belong in Maylandia, they only suggest its inclusion, they don’t actually state it 

is to be included. 

The other “evidence” for Maylandia 

By now you are probably quite confused, which is not surprising as MEYER & 

FOERSTER’s work is itself totally confused, and it is essential that scientific work 

should be clear and unambiguous. But I hope that you also now understand why 

Maylandia does not satisfy Article 13 (wording and intention) and hence is 

unavailable, a nomen nudum with no differentiating characters, not even for the 

zebra complex. 

Nevertheless I think it is necessary to also “poke holes” in some of the other 

“evidence” adduced by Maylandia supporters discussed by SCHRAML. 

42 BAILEY: the case for Metriaclima

KULLANDER (1999): laudably - and rather over-emotionally - defends the Principle 

of Priority, which states that the oldest name is that which should be used, but 

omits to mention that the Principle of Priority applies only to names that satisfy 

the criteria of availability (for our purposes = validity) laid down by the Code. He 

adduces no arguments to defend the availability of Maylandia, and it is not 

enough simply to say a name is available in disputes like this. Even if you are a 

world-famous cichlid taxonomist you must back up your statements with 

arguments and facts. So strike KULLANDER from the debate. 

CONDÉ & GÉRY (1999) assert that Article 13d of the 1985 edition of the Code 

applies, and hence Maylandia is valid as MEYER & FOERSTER described the 

subgenus and its type species in a single description. However, Article 13d 

applies only to monotypic genera (those containing one species, where there is 

no point in separate descriptions as the genus is based on just the one species). 

But we have another school of thought arguing that Maylandia is valid because 

MEYER & FOERSTER provide a description by listing a set of characters that apply 

to the Pseudotropheus zebra complex which also belongs to the new subgenus. If 

you subscribe to that view then this means that in writing their paper MEYER & 

FOERSTER were aware that Maylandia was not monotypic and so Article 13d 

cannot possibly apply. Moreover, as SCHRAML points out, the rule doesn’t apply to 

subgenera (though why is a mystery, but the Code admits to not being perfect, so 

it could be an oversight). So strike CONDÉ & GÉRY ………….! 

KONINGS (2005) made some small errors in his translation of MEYER & FOERSTER. 

So? These were of no relevance whatsoever to his arguments. Perhaps the object 

of pointing out these errors was to discredit KONINGS’ understanding of the paper 

in general? I can see no flaws in KONINGS’ arguments based on his understanding 

of the paper. 

The abstract to MEYER & FOERSTER’s paper states that they are describing a 

species and subgenus. So? Abstracts are merely brief summaries designed to tell 

the reader what a paper is about, so he can decide whether to read it. They are 

of dubious taxonomic validity as they are often written by the publisher, and 

where they appear in more than one language they frequently do not say 

precisely the same thing. MEYER & FOERSTER’s paper contains abstracts in 

English and German (but not in the language of the paper – French – as one 

would expect), and these two abstracts are so very different that they might be 

summarising two different papers! So, which abstract do we take as cogent and 

valid? The English one cited by SCHRAML, or the German one (MEYER & FOERSTER’s 

mother tongue) which does not make the statement about the subgenus and 

species? Search me! 

One argument not mentioned by SCHRAML, but which I’ve seen elsewhere, is that 

Dr ETHELWYNN TREWAVAS peer-reviewed the paper, as stated by its authors. That 

may be true, but it doesn’t mean that she approved it, for all we know she may 

have said it was utter rubbish. Her own paper published in the same issue of the 

journal is, after all, to infinitely higher taxonomic standards and has not, as far 

as I know, been challenged. There is no obligation on authors and editors to take 

any notice of reviewer comments, though editors usually do and have the power 

to reject papers unless amended or justified by the author. I would mention that 


43

peer review is supposed to be anonymous, and the fact that in this case it clearly 

was not suggests to me that the journal’s peer-review policy was at best not as 

strict as desirable. 

Retro-active or not? 

SCHRAML asks whether the setting of a date from which a new rule applies is truly 

intended to be retro-active, ie affecting names previously regarded as valid 

under the previous Code. The answer is, quite simply, YES! The introduction to 

the 1985 Code makes this quite clear: “(6) The Code provides guidance for 

zoologists needing to establish new names, and rules to determine whether 

any name, previously proposed, is available and with what priority; whether 

the name requires amendment for its correct use……” (my bold type). From 

this it is quite clear that the Code is intended to be retro-active. 

The latest (2004) edition of the Code caused some major retroactive changes to 

names, most notably the proper way to deal with umlauted u (ü) in names based 

on proper names, such that u or ue is now correct depending on a cut-off date, 

and names after that date need to be corrected where they do not conform. Even 

more far-reaching is the requirement by the ICZN for the dates of names to be 

the date when the paper appeared in print. In days of yore, papers, for example 

those of the Zoological Society of London, were frequently read to a meeting and 

only printed later, and the date of the reading was taken as the date of the paper. 

Not any more! It now has to be the ACTUAL date of publication, and this has 

caused huge confusion. For example, REGAN’s revision of Lake Malawi cichlids, 

cited in much of the literature and a host of names as REGAN, 1921, is now REGAN, 

1922. 

SCHRAML has questioned the relationship between these retro-active rules and 

stability in nomenclature, one of the most important objectives of the Code. But 

in fact it’s quite simple. Although the Code is designed to be absolutely explicit 

and unambiguous in its wording in order that the INTENTION of the rule is 

fulfilled, sometimes this objective is not achieved and clarification of a rule is 

required, by rewording, and where the old, inadequate rule has caused problems, 

then to put these right the new rule must be retro-active. Nowadays we write 

specific names with a lower case letter – greshakei – and hyphens aren’t allowed, 

but it wasn’t always thus, many names have been corrected retro-actively, even 

before the first edition of the Code was published in 1961. Was that wrong too? 

We should, perhaps, consider that the fundamental date of 1st January 1758, 

from which modern taxonomy is taken to begin, was not selected at that time but 

much later, and the decision taken that – retroactively – all names dating from 

before that date would be disregarded. 

Fall-out 

There has been a certain amount of unfortunate “fall-out” linked to the dispute, 

in part a function of our electronic age. I have been told by a number of authors, 

who believe Metriaclima to be the correct name, that they have been obliged to 

use Maylandia in order to have their work published, as the editors of some 

44 BAILEY: the case for Metriaclima

scientific journals and some peer-reviewers are stating that Metriaclima is 

invalid because Fishbase or some noted taxonomist has plumped for Maylandia. 

One thing must be made clear – no organisation such as Fishbase, no institution, 

and no individual such as an editor or peer-reviewer has the right to pre-empt the 

role of the ICZN in ruling on nomenclatural disputes. I can only deplore such 

attempts to impose a personal, editorial, or institutional viewpoint on the 

freedom of every author to decide for him- or herself which name to follow until 

a ruling is made by the ICZN. 

In this regard I cannot commend too highly the editorial policy of eggspots in 

allowing authors their rightful freedom of taxonomic expression. 

References: 

CONDÉ, B. &. GÉRY, J. (1999): Maylandia MEYER ET FOERSTER, 1984, un nom générique disponible (Teleostei, Perciformes, Cichlidae). 

Revue française d'Aquariologie Herpetologie, 26 (1-2): 21-22. 

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE (ICZN) (1985): International Code of Zoological Nomenclature, 3rd edition. 

INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE (ICZN) (2004): International Code of Zoological Nomenclature, 4th edition. 

KONINGS, A. (2005): Maylandia or Metriaclima, ...again! The Cichlid Room Companion: http://www.cichlidae.com/article.php?id=355 

KONINGS, A. F. & STAUFFER, J.R. JR. (2006): Revised diagnosis of Metriaclima with description of a new species (Teleostei: Cichlidae) from 

Lake Malawi National Park, Africa. Ichthyological Exploration of Freshwaters, 17 (3): 233-246. 

KULLANDER, S. O. (1999): Diskussionsbeitrag für Cichlid-L. 

MEYER, M. K. & FOERSTER, W. (1984): Un nouveau Pseudotropheus du lac Malawi avec des remarques sur le complexe Pseudotropheus- 

Melanochromis (Pisces, Perciformes, Cichlidae). Revue française d'Aquariologie Herpetologie, 10 (4) [1983]: 107-112. 

SCHRAML, E. (2009): Maylandia oder Metriaclima - immer noch nicht geklärt? eggspots, 2: 5-15. 

STAUFFER, J. R., JR. , BOWERS, N. J., KELLOGG, K. A. & MCKAYE, K. R. (1997): A revision of the blue-black Pseudotropheus zebra 

(Teleostei: Cichlidae) complex from Lake Malawi, Africa, with a description of a new genus and ten new species. Proceedings of the 

Academy of Natural Sciences of Philadelphia, 148: 189-230. 

TREWAVAS, E. (1984): Nouvel examen des genres et sous-genres du complexe Pseudotropheus-Melanochromis du lac Malawi. Revue 

française d'Aquariologie Herpetologie, 10 (4): 97-106. 


45

Eggspots Elsewhere 

Articles on cichlids with eggspots elsewhere in the recent literature 

Period February - September 2010 

Amazonas (http://www.ms-verlag.de/AMAZONAS.121.0.html) 

6 (5), No. 31, September/October 2010: 

- SZILLAT, K.: Arterhaltung im Wohnzimmer. (Yssichromis piceatus) Pp. 58-59. 

AqualogNews (www.animalbook.de) 

No. 94: 

- SCHÄFER, F.: Placidochromis cf. phenochilus "Tanzania" - nicht nur schön, sondern auch 

friedlich. Pp. 24-25. 

Aquaristik Aktuelle Süßwasserpraxis (http://www.aquaristikonline.de/index_aquaristik.html) 

18 (5): 

- STAECK, W.: Buntbarsche aus dem Viktoriasee. Oft attraktiv - doch selten gepflegt. Pp. 40- 

45. 

Aquaristik Fachmagazin (www.tetra-verlag.de/index.htm ) 

42 (1), No. 211, February/March 2010: 

- SEIDEL, I.: Eine Zuchtfarm für Malawi-Buntbarsche in Florida. Pp. 12-16. 

Buntbarsche Bulletin (Journal of the American Cichlid Association, 

www.cichlid.org/index.php?pageid=buntbarsche_bulletin) 

No 256, February 2010 

- STEEVES, G.: Species at Risk Highlights. ('Haplochromis' thereuterion, Xystichromis sp. 

'Kyoga Flameback', 'Haplochromis' sp. 'fine bar scraper'). P. 3. 

No 258, June 2010 

- STEEVES, G.: Species at Risk Highlights. (Astatotilapia desfontainii, Lipochromis sp. 

"Matumbi Hunter", Prognathochromis perrieri, Pyxichromis orthostoma). P. 3. 

- STEEVES, G.: Pyxichromis orthostoma. P. 25. 

Cichlidae (Nederlandse Vereniging van Cichlidenliefhebbers NVC; 

www.nvcweb.nl/portal/) 

36 (2) 2010: 

- VAN HEUSDEN, H.: Vijf Orthochromis-Soorten van Tanzania. Pp. 5-29. 

36 (3) 2010: 

- KONINGS, A.: De Mbuna van het elongatus-Complex, Malawimeer. Pp. 5-13. 

Cichlid News (Aquatic Promotions Inc., Miami, USA, www.cichlidnews.com ) 

19 (2) April 2010: 

- KONINGS, A.: Sex determination and the OB pattern in Malawi cichlids. Pp. 6-13. 

46 Eggspots Elsewhere

19 (3) 2010: 

- VAN HEUSDEN, H.: Orthochromis from Tanzania: Report of a Collecting Trip to the 

Malagarasi Basin – Part 1. Pp. 6-14. 

- KONINGS, A.: The Largest Cichlid of Lake Malawi. Pp. 15-19. 

19 (4) October 2010: 

- KONINGS, A.: Pseudotropheus elegans rediscovered. Pp. 14-17. 

- VAN HEUSDEN, H.: Orthochromis from Tanzania: Report of a Collecting Trip to the 

Malagarasi Basin – Part 2. Pp 20-27. 

- ANDERSEN, T.: Observations on Limnochromis staneri POLL, 1949. Pp 6-12. 

DATZ Die Aquarien- und Terrarienzeitschrift (Ulmer Verlag; ISSN 1616- 

3222; www.datz.de/) 

63 (8) 2010: 

- SCHRAML, E.: Der Kleine oder Vielfarbige Maulbrüter. Pp. 4-6 in Aquarien-Praxis. 

DCG-Informationen (Deutsche Cichliden Gesellschaft e.V.; ISSN 0724-7435; 

www.dcg-online.de/noFrames/files/index.html) 

41 (3) 2010: 

- LOOSE, S.: Unterwegs am Nordende des Tanganjikasees. 1. Teil: Burundi 2009. Pp. 50- 

67. 

41 (4) 2010: 

- SCHRAML, E.: Taeniolethrinops sp. "Black Fin Pombo". Pp. 81-84. 

41 (6) 2010: 

- STAECK, W.: Zur Entstehung zwischenartlicher Kreuzungen von Cichliden im 

Gesellschaftsaquarium. 1. Teil. Pp. 139-146. 

41 (7) 2010: 

- STAECK, W.: Zur Entstehung zwischenartlicher Kreuzungen von Cichliden im 

Gesellschaftsaquarium. 2. Teil. Pp. 154-162. 

- BAUER, R.: Iodotropheus sprengerae OLIVER & LOISELLE, 1972. Ein friedlicher Cichlide für 

kleinere Aquarien? Pp. 171-176. 

41 (9) 2010: 

- MORGENSTERN, R.: Astatotilapia calliptera (GÜNTHER, 1893). Der "andere" Malawisee- 

Cichlide. 1. Teil. Pp. 202-211. 

Practical Fishkeeping (PFK) (www.practicalfishkeeping.co.uk) 

Aug (8) 2010: 

- KONINGS, A.: Ad Konings introduces the kambuzi cichlids of Lake Malawi. Pp. 48-50. 


47

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now we would like to request a voluntary contribution towards defraying our costs. 

Hitherto we have charged 1.50 Euros per issue, and we would be most grateful if you would 

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48 

Book review: AXEL BÖHNER (2010): Die 

Viktoriasee-Cichliden in Deutschland. Ein 

Leitfaden für die Pflege der Buntbarsche aus 

dem Viktoriasee. Books on Demand, 

Norderstedt. ISBN 978-3-8391-3286-9, 26.90 €. 

In the 146 pages of this book AXEL BÖHNER has 

managed to cover, in words and photos, almost all 

the cichlid species of the Haplochromis 

assemblage currently available in Germany, 

totalling more than 60 in number. For all of them 

the author provides a summary of distribution, 

size, coloration, the appropriate aquarium, 

feeding, social behaviour, breeding, and so on. 

When it comes to scientific names BÖHNER 

conforms with GREENWOOD’s proposals, but at the 

same time he doesn’t omit to include synonyms 

and trade names. The names used are also correct, 

with a few exceptions (for example Astatotilapia 

nubila). The species are all illustrated with colour 

photos. It is true that some are rather small 

(though several views of the species are generally given to compensate) and because 

of the paper quality the print standard isn’t quite up to that usual for glossy 

magazines, but nevertheless the overall impression can be described as generally 

attractive. The book is rounded off by an introductory section discussing Lake Victoria 

and its problems, the habitats of the cichlids in the lake, and general details of 

maintenance and breeding, creating a successful reference work, written by an 

enthusiast for enthusiasts. Published in German only.

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