A glimpse of the complexity of the genus Tropheus.

This article is a compilation of the seven entries from the "Tropheus Week" published March 16-22, 2015. During the week a new entry on Tropheus was posted daily on African Diving's social media. The article contains some additional material in form of pictures and map plus a few comments by the authors in response to questions from readers. Also, the Day 6 entry has been translated into French and is available via a link in the end of the same paragraph.

Fig. 1. The "Tropheus Week" primarily addressed seven lesser known Tropheus variants.

Day 1 - Tropheus sp. “Limespot”: Poll’s and Brichard’s observations on Moore’s Tropheus (part 1 of 2)
Tropheus moorii is one of the most popular species of Lake Tanganyika cichlids among hobbyists and scientists alike. It was described by Boulenger in 1898 based on specimens collected by Moore in the southernmost part of the lake. Subsequent discoveries of additional Tropheus populations were usually referred to as geographical colour variants of T. moorii. This was also applied by Poll (1956: 100f) when describing the live colouration of several Tropheus populations from the Congo, referring to them as T. moorii. Some of Poll’s material originated from around Tembwe Bay and Mwerazi. Specimens from these localities, however, had a meristic anomaly. Instead of possessing six anal-fin spines, as in Boulenger’s specimens of T. moorii, Poll’s specimens from Tembwe Bay and Mwerazi had only five spines. They were, nevertheless, referred to as T. moorii. Poll (1956: 100; Pl. IV, fig. 3) described the live colouration of the Mwerazi variant as being “olive green with a yellowish belly”, and having “spots and irregular stripes of a beautiful lemon yellow colour, on the flanks of the body”, and in addition, “the dorsal and the caudal fins are adorned with lemon-yellow spots”. Following Poll in referring the Mwerazi variant to T. moorii, Scheuermann (1976) and Staeck (1985) also described it as being olive green with lemon-yellow spots and stripes, as well as having a yellowish belly.
In June 1984 Pierre Brichard discovered several new Tropheus moorii-like populations between Moba and Kapampa in the Congo, including the already known variant from Mwerazi. The latter was described as being “emerald-green, with a large lemon-coloured patch” behind the pectoral fins (Brichard 1989: 177). Other variants from neighbouring localities were also described as exhibiting such “a lemon patch on the side” or as having a “conspicuous broad circular patch 3 to 4 cm across just behind the pectoral fin. This patch is a pale yellow, pale green, or nearly white” (1989: 177). Brichard’s impression of this group of populations was that it “appears to be basically different from any we have seen in the lake so far”, alluding to the contrasting colouration. The most specific difference from similar Tropheus species in this part of the lake is the number of anal-fin spines: having five spines instead of six. Partially based on this character, Brichard classified this group of populations as a distinct lineage (here interpreted as a distinct species). This meristic character (the number of anal-fin spines) has always been important in Tropheus classification, and was initially the primary diagnostic character when separating the southernmost Tropheus (T. moorii Boulenger) from the northernmost (T. sp. “Black” Konings) (Nelissen 1979) {see a discussion on the anal-fin spines in the blog article about ‘Brichard’s Tropheushere:}.
All Tropheus variants from the Moba – Kapampa vicinity are commonly identified as T. moorii. However, following Brichard’s discoveries and observations, this is not a correct identification, and they are here referred to as T. sp. “Limespot”. Brichard’s documentation of this species (T. sp. “Limespot”) only concerned the geographical variants he found along the Congolese coast, but the similar variants of this species present along the Tanzanian coast (Lyamembe – Kalya) diagonally opposite to the northernmost part of the mentioned Congolese coast (Moba – Kapampa) are near identical, with no obvious specific differences, and therefore likely conspecific.

Fig. 2. Tropheus sp. "Limespot" at Lyamembe, depth 5 metres. This species is found from around Mabilibili Bay to the northern side of Nanga Bay. Further south along the rocky shores at Nanga Point, Makola and Sibwesa Point (see map Fig. 10) the species is absent, until at Kalya village where it reoccurs at the shallow reefs along the shore.

Day 2 - Tropheus sp. “Limespot”: Conspecific populations in the Congo and Tanzania (part 2 of 2)
The name Tropheus sp. “Limespot”, as this species is referred to here, dates back to both Poll’s description of the Mwerazi variant as having “spots and irregular stripes of a beautiful lemon yellow colour” and “lemon-yellow spots” (1956: 100), and to Brichard’s description of the species, to which the name is perhaps more significant: “Behind the pectoral fin extends a very large circular patch in strong contrast with the surrounding area. This patch is either lemon yellow or a ghostly white and is the only contrasting feature of the colour pattern” (1989: 176).
The specific character of Tropheus sp. “Limespot” distinguishing it from T. moorii is the number of anal-fin spines: five in T. sp. “Limespot” and six in T. moorii. The “striking colour pattern” as mentioned by Brichard is only vaguely, if at all, separable from T. moorii, which also goes for the tail fin, which is said to be larger. However, it is now possible with more advanced technology to detect even minor colourational and shape-wise differences among Tropheus populations (Herler et al. 2010). Further dissimilarities may be exposed, pending a thorough analysis.
The geographical distribution of T. sp. “Limespot” in Tanzania (Lyamembe – Kalya), which is patchy due to the sub-optimal habitat, covers about 20 km. The variant of T. sp. “Limespot” found along the rocky shore just north of Sibwesa was occasionally collected and exported by us in the nineties, by the name of T. moorii “Kibwesa” (Zadenius 1995: 43). The variant of T. sp. “Limespot” found at Nanga (in between Lyamembe and Sibwesa) was sampled by us for scientific studies in 2007 and deposited at the Swedish Museum of Natural History as Tropheus sp. “Limespot – Nanga”.
The Congolese variants of Tropheus sp. “Limespot” were collected and exported by African Diving for the first time in 1996 and concerned primarily the following four variants:

- Tropheus moorii “White limespot – Mwerazi”
- Tropheus moorii “Limespot yellow – Kapampa”
- Tropheus moorii “True limespot – Masanza”
- Tropheus moorii “Sunspot – Longola”

At the end of 2006 we collected Tropheus sp. “Limespot” at Kalya; totally 40 males and 84 females. Among the males were 2 individuals with 4 anal-fin spines, 24 individuals with 5 spines, and 14 individuals with 6 spines. Among the females were 8 individuals with 4 anal-fin spines, 52 individuals with 5 spines, and 24 individuals with 6 spines. The four spines may suggest some kind of hybridisation with T. polli, which has four anal-fin spines in this area. However, the presence of six anal-fin spines is something of a mystery, if this too is to be considered as a result of hybridisation. There are no Tropheus species in the vicinity of Kalya with exclusively six anal-fin spines. The closest northwards is T. sp. “Lukuga” at Lufungu Bay, 40 kilometres away {see blog article on ‘Brichard’s Tropheushere}, and southwards is T. sp. “Mpimbwe” {see the variant “Yellow cheek” here and a map of Cape Mpimbwe here}. In summary, roughly two thirds of the individuals of T. sp. “Limespot” at Kalya correspond to the species characteristic of having five anal-fin spines, while one third does not. However, the latter is for the time being not interpreted as being non-conspecific, but rather as a possible result of hybridisation. (N.B. The fishes were later on, during 2007, exported to Åleds Akvarium in Sweden {link here}.)
Tropheus sp. “Limespot” largely corresponds to ‘Tropheus of lineage 6’ in Schupke (2003). The latter is a lineage, or species, which is stated to comprise a peculiar colouration and “a striking stripe pattern on the forehead” (2003: 69). T. sp. “Limespot” is primarily characterised on the number of anal-fin spines, which is an important diagnostic character in Tropheus, besides colouration. In contrast to ‘Lineage 6’ (Schupke 2003), T. sp. “Limespot” includes all Congolese Tropheus populations between Moba – Kapampa, because they all seem to possess an anal fin of five spines (Brichard 1989: 152, table; 176), as opposed to six spines in T. moorii (Boulenger 1898).
The molecular analysis and the phylogenetic inference of Baric et al. (2003) identified groupings of Tropheus. One of the DNA-based lineages was the so-called ‘Lineage D’, which was seemingly conforming to the morphology-based identification of T. sp. “Limespot”. The analysis also confirmed the presence of T. sp. “Limespot” populations on opposite shores — in both the Congo and in Tanzania. Evidently, the molecular analysis reflects the existence of T. sp. “Limespot” as it is presented here.

Fig. 3. Tropheus sp. "Limepsot", brooding female at Kalya, depth 4 metres. The habitat at Kalya consists of submerged sandstone reefs running along the shoreline.

Day 3 - Tropheus “Golden Kushangaza”: Colourational anomaly at Udachi Point in Lake Tanganyika
On the 4th of April 1990 our team went diving at Udachi Point. This is a rocky habitat located just a few hundred metres from our camp at Katoba, near Udachi {see map here}. One of the divers, Yusuph Maridadi, made a truly remarkable discovery by spotting a conspicuous golden coloured Tropheus among the regular black individuals. This first golden individual made its way to the cover of the Journal of the Nordic Cichlid Association the same year (African Diving Ltd 1990; Karlsson and Lundblad 1990) {see the cover here}. This was the first ever documented case of a mutant coloured Tropheus. The name “Kushangaza”, meaning “surprisingly” was applied to the fish. A total of 11 Tropheus “Golden Kushangaza” individuals were collected in 1991 and sent to Sweden for breeding purposes (Zadenius 1991). Because of the scarcity of the fish, very little collecting efforts were made over many years to come.

Fig. 4. Kushangaza form of Tropheus sp. “Mpimbwe” (“Golden Kushangaza”) at Udachi Point in 4 metres depth. Discovered in 1990, the Udachi Kushangaza was the first ever documented case of mutant coloured Tropheus {see video sequences of three different individuals; sequence 1, sequence 2, sequence 3}.

Day 4 - Natural hybridisation in Lake Tanganyika: Tropheus “Red blotch” (part 1 of 2)
Apart from being taxonomic units, species are among traditional biologists (and especially among proponents of the ‘biological species concept’) also strongly argued to be ‘real entities’ with special species-level qualities (such as ‘reproductive isolation’), as opposed to "poorly differentiated way-stations in a continuous hierarchy of biodiversity" (Mallet 2005). Hybridisation between species is the opposite of reproductive isolation and challenges the ‘real entities’ in biological species. In general, there is a strong reluctance to acknowledge hybridisation as a fairly common occurrence in nature. However, it is thought that at least 10% of animal species are involved in hybridisation, and species in nature are often incompletely isolated (species are in gene-flow contact) for millions of years after their formation (Mallet 2005).

There are at least two localities along the Tanzanian shores of Lake Tanganyika where one may find natural hybridisation between two sympatric (or partially fused) Tropheus species. One of these localities is in the Mabilibili – Lyamembe vicinity, where individuals of mixed origin are fairly common. Based on both underwater observations and aquarium studies, we have over the years (especially in recent years) identified about 40 fairly distinct colour morphs which are frequently occurring among these hybrid individuals. These many distinct hybrid morphs are possible to identify by obtaining details on a particular set of characters and the state they are in, e.g. dorsal fin may be red, yellow, or blue; cheek colour may be red, yellow or black, etc.

Fig. 5. A natural hybrid individual photographed in 3 metres depth at Lyamembe. The hybridisation involves Tropheus sp. "Limespot" and Tropheus sp. "Doubleblot". In the Mabilibili/Lyamembe area we have identified and documented (pictures) about 40 fairly distinct hybrid colour morphs, ranging from almost pure Tropheus sp. "Limespot" to almost pure Tropheus sp. "Doubleblot".

Along the rocky shores of Mabilibili and Lyamembe there are three sympatric Tropheus species: T. polli, T. sp. “Doubleblot/Kirschfleck”, and T. sp. “Limespot” {see above Day 1 and Day 2}. In the same habitat also the hybrids are found. The latter is highly likely the result of hybridisation between any two of the genuine species; perhaps, the hybridisation may even involve all three species. However, we have since 1994 {see post tomorrow} and onwards observed (in the lake and in aquaria) altogether about 350 individuals, and none of these have possessed a lunar-shaped caudal fin, or a forked caudal fin with elongated tips, as is possessed by T. polli in this locality. Therefore, the hybridisation is most likely only between T. sp. “Doubleblot” and T. sp. “Limespot”. Of course, hybrid individuals may also take part in the reproduction, and not only ‘pure’ individuals from either of the two species. The result is a multitude of phenotypes, with a ‘successive transition’ between the colouration of the two species.
Hybrid individuals from the Mabilibili – Lyamembe vicinity were first exported by us in 1995 (Åleds Akvarium Sweden, March 5, 1995). The particular hybrid morph being mostly red (belly, flank of body, and cheek; see picture in this post) was labelled T. “Red blotch” in at least one shipment to the same importer. Furthermore, another hybrid morph similar to the phenotype of the genuine T. sp. “Doubleblot”, but with larger and irregular blots, together forming a single large blot, as well as being even more red than the genuine T. sp. “Doubleblot”, was exported to Sweden in December 1994, and reported as T. brichardi “Double blot Kibwesa” (Zadenius 1995). Four hybrid individuals of the morph T. “Red blotch” were collected by us in 2007 for scientific studies, and later deposited at the Swedish Museum of Natural History as T. “Lyamembe”.
Moreover, already in 1991 we discovered and exported a new interesting species of Tropheus which we named T. sp. “Red belly” (Zadenius 1991, 1992). This is, by all accounts, a genuine species with a characteristic which seems to be out of the ordinary. T. sp. “Red belly” does not originate from the Mabilibili – Lyamembe vicinity. We will report in more detail on this species in a future article.
In Schupke (2003) and Egger et al. (2007), however, the name T. “Red belly” appeared as the name of the hybrid individuals from the Mabilibili – Lyamembe area. This is an unfortunate coincidence, or perhaps rather a misconception and failure by alternative or succeeding collectors in communicating the original names. The name will then pertain to two different ‘species’; one may call it a homonym — the same name applied to two different taxa. We will continue to use the name T. sp. “Red belly” for the genuine species found elsewhere, and T. “Red blotch” for the ‘red blotched’ hybrid individuals found in the Mabilibili – Lyamembe vicinity.

Fig. 6. Tropheus “Red blotch” at Mabilibili, depth 5 metres. This hybrid individual is the result of hybridisation between T. sp. “Doubleblot” and T. “Limespot”. Reddish hybrid individuals at Mabilibili/Lyamembe area should not be confused with Tropheus sp. "Red belly" which is a different species that originates elsewhere. The latter was discovered and named by our team in 1991.
In 2007 we collected two brooding females of the “Red blotch” morph (later preserved) at Lyamembe. Each brood was subsequently kept separated in aquaria until reaching adult size when several hybrid morphs including a few almost pure T. sp. “Limespot” and T. sp. “Doubleblot” were revealed among the individuals. None of these developed a lunar-shaped caudal fin with elongated tips. Such a characteristic feature is typically seen in T. polli, another species which is co-existing with the two former species in this area. Thus, T. polli is unlikely included in this hybridisation event. Moreover, no individuals of the latter diverging from the phenotype have been observed in the area.

Day 5 - Natural hybridisation in Lake Tanganyika: Tropheus “Zadeni” (part 2 of 2)
Is hybridisation a reproductive mistake?
Most hybridisation occurs between closely related species, and the progenies are in such cases not necessarily less viable or sterile. The hybrid origin of species has been discovered by using molecular genetic techniques, suggesting that many more species are of hybrid origin than was previously thought (Seehausen 2004). Moreover, it seems that hybrid species may become isolated by other means than by ‘reproductive isolation’. Hybridisation in nature may be an evolutionary impulse providing genetic diversity and faster-evolving adaptive phenotypes in a changing environment. At least one cichlid species in Lake Tanganyika (Neolamprologus marunguensis) is suggested being of a hybrid origin (Salzburger et al. 2002).
Within the ‘modern haplochromines’, which include Tropheus, non-hybridisation events between closely related species are often maintained by colour-assortative mating. In addition, sexual selection has been suggested as having contributed to much of the evolution of the many geographical colour morphs distributed along the length of the lake. In Tanzania alone, at least 50 fairly distinct geographical colour variants of Tropheus may be identified, which are confined to a particular locality (pers. obs.). However, not all sympatric Tropheus species seem to maintain the colour-assortative reproduction, even when two majorly different colour patterns are involved.

Fig. 7. Tropheus sp. "Doubleblot" (also known as "Kirschfleck") at Lyamembe, depth 5 metres. Lyamembe Point is the southernmost distribution of the species. In the Mabilibili/Lyamembe area where the species is sympatric with Tropheus sp. "Limespot", the two species hybridise.
Fig. 8. Young brooding female of Tropheus polli at Lyamembe in 8 metres depth. Even though T. polli occurs in the same area as T. sp. "Limespot" and T. sp. "Doubleblot, this species does not take part in the hybridisation and is phenotypically homogeneous in this locality.

In the Mabilibili – Lyamembe vicinity two species are frequently hybridising. These are T. sp. “Doubleblot/Kirschfleck” and T. sp. “Limespot”. The result is a large number of hybrid colour morphs, each of which resembles a colourational mixture of the two 'genuine' species. We have since 1994 {see above, Day 4} and onwards observed in the Mabilibili – Lyamembe vicinity altogether about 350 hybrid individuals. None of these have possessed a lunar-shaped caudal fin as is possessed by T. polli in this locality. Thus, the hybridisation is unlikely to include the latter species, but only T. sp. "Doubleblot" and T. sp. "Limespot". About 40 distinct hybrid morphs are identified and occur in the area (Mabilibili – Lyamembe), of which T. “Red blotch” may be the most conspicuous {see above, Day 4 paragraph}.

Another beautiful hybrid colour morph from this locality is Tropheus “Zadeni” (see Fig. 9). This morph was discovered in 1994 and exported to Åleds Akvarium in Sweden on March 5, 1995. The shipment comprised four individuals (two males and two females) and they were labelled Tropheus moorii “Zadeni”. The latter name was in honour of Marcus Zadenius, one of Sweden’s earlier Tropheus enthusiasts. The four T. “Zadeni” individuals were later supposedly deposited at the Museum of Zoology of Lund, in southern Sweden, for scientific studies. Tropheus “Zadeni” remained a much sought-after Tropheus variant (Zadenius 1997). To this very day (March 20, 2015) the collecting site for Tropheus “Zadeni” has been kept a secret.

Fig. 9. Tropheus “Zadeni” at Mabilibili, depth 5 metres. This hybrid individual is the result of the hybridisation between T. sp. “Doubleblot” and T. “Limespot”. The "Zadeni" hybrid morph was exported to Åleds Akvarium in Sweden on March 5, 1995 by the name Tropheus moorii "Zadeni".

Comments by the authors on hybridisation in responce to questions from readers

Comment 1: This is natural hybridisation without the interference of mankind. The hybridisation has likely been going on for about 18000-12000 years. Lake Tanganyika has in the past repeatedly been divided into at least three large parts, so-called proto-lakes. The time period of 18K-12K years is how long ago it was since the recent central proto-lake (originally inhabited by only one of the two species) and the southern proto-lake (likely inhabited by the other species) met and fused into one bigger lake, which was due to a rising lake level (for ref. re. lake-levels, see e.g. Sturmbauer et al. 2005; ref. list here). At this point the evolutionary paths of the two species crossed. Why the two species/populations have not already long ago fused into one is subject for the viability in hybridisation and may be dealt with in a future article.

Comment 2: At Mabilibili-Lyamembe, where the distribution of T. sp. "Doubleblot" and T. sp. "Limespot" overlap, the two sympatric species partially hybridise. The colouration of the hybrid individuals seems to be a mixture of the colouration of the two species. But the hybrid colouration is not uniform; it is not fixed to a particular colouration, or phenotype, as it is in the two 'genuine' species. It varies between individuals. Some of the hybrids are more reddish without stripes (resembling T. sp. "Doubleblot"), others are more greenish brown, and striped (resembling T. sp. "Limespot"), and yet others are a 'perfect mix' resembling both species just as much (or rather, just as little; e.g. T. "Zadeni").

We have identified about 40 fairly distinct hybrid colour morphs. Largely all of the hybrid individuals can be sorted to any of these 40 colour morphs.

For the difference between the hybrid morphs "Red blotch" and "Zadeni", see the pictures; the former is only red/reddish, not yellow, and without stripes; the latter is both reddish and yellowish, along with stripes, and so on.

T. sp. "Red belly" is a different species. It does not exist in the area of Mabilibili-Lyamembe. We will report in more detail on this species in a future article.

Fig. 10. Central parts of Lake Tanganyika with Lyamembe area enlarged. The map is showing distribution range for Tropheus sp. "Limespot, Tropheus sp. "Doubleblot/Kirschfleck", and the area where natural hybrids between the two species occur.
Fig. 11. The rocky outcrop at the southern side of Mabilibili Bay. The cement beacon in the picture to the right marks the southern boundary of Mahale Mountains National Park (also see map Fig. 10). Fishing within the borders of the national park is strictly forbidden. More on Mahale Mountains NP here).

Day 6 - Tropheus moorii “Murago Tanzania”: On the fast track to extinction?
The small population of Tropheus moorii “Murago Tanzania”, discovered in 2008, was found by commercial collectors in 2014, when the collecting of this Tropheus variant commenced. Announcements addressing the fragile nature of this small population were published on several occasions to inform importers and hobbyists alike (Karlsson 2012, 2014a,b; Konings 2014).

Sadly, it seems that collections continue of T. moorii “Murago Tanzania”, and exports are repeatedly containing the fish. We fear that the population is already seriously diminished and may well have met a viable point by now, with the risk of the loss of genetic variability due to the low number of individuals remaining. With less genetic variability, the population may lose its virility, reduced fertility, and become more vulnerable to genetically related problems. In the long term, a small population with less genetic variability is less likely to respond to selective pressure and more likely to become extinct. In the short term, a small population will be more vulnerable to diseases and parasites than populations with a greater gene variation. This applies at the individual level as well, because with fewer individuals the risk of inbreeding increases, with the result being a higher number of individuals potentially having two defective copies of a particular gene.
The risk of literally wiping out the entire Tropheus population is also imminent by the collecting of the major part of the population, leaving the remaining individuals to harsh competition by other species, predators, and with major difficulties to find a mate. Obviously, there are no reasons to discuss the aspect of inbreeding and genetic loss when the whole population is gone.
Several authors (Ad Konings, Livio Leoni, Benoît Jonas, Hans-Georg Evers and Wolfgang Staeck) have contributed to the conservation of this unique and highly threatened Tropheus variant (see reference list). Also importers are taking a stand against the trade with wild “Murago Tanzania” (Steve Ewald and others).
A very fresh report from an individual Tanzanian collector says that it now seems that most adult individuals of “Murago Tanzania” are gone from the habitat. There are at least six ‘companies’ collecting “Murago Tanzania”. When collecting, the entire small habitat is repeatedly swept over while searching for the fish. Collectors are now collecting even on the fringes of the area of distribution, where the fish is extremely scares due to suboptimal habitat and where the pattern of colours, stripes and spots are increasingly faint.
Please do not purchase wild Tropheus moorii “Murago Tanzania”. Bred fish from already imported wild individuals or fish bred at the lake should soon be (and perhaps already are) available from breeders/dealers.
Let us hope that the report from the Tanzanian collector is not entirely correct, and that there still are enough individuals (sub-adults and juveniles) in the habitat for the population to be able to recover from recent collections, though we fear that the population of "Murago Tanzania" may already have reached a minimum viable population size.
“The rates of extinction today far surpass those of the most apocalyptic mass extinctions our planet has ever experienced.” (Ward 2004)
Fig. 12. Tropheus moorii "Murago Tanzania", female in its natural habitat, depth 5 metres. This conspicuous geographical variant was discovered in May 2008.

Day 7 - The ‘Green Wimple’ of Lake Tanganyika: Tropheus sp. “Crescentic”
Tropheus sp. “Crescentic” was first identified by Pierre Brichard based on populations he observed in the Congo. In addition, conspecific populations also occur in Tanzania, between Malagarasi River and Halembe. The most conspicuous feature may be its crescentic tail fin. Some of the variants from around Karago and Cape Kabogo have earlier been referred to as ‘Green Wimple Tropheus’, alluding to the pennant-shaped tail fin. The variant with the yellow pectoral spot was for a long time thought to originate from Isonga, north of Ikola. However, our team rediscovered this yellow-stained variant in 1992 along the rocky coast between Mkuyu and Segunga, not very far from Cape Kabogo. Evidence of the existence of T. sp. “Crescentic” has also been found in several molecular phylogenetic analyses.
Read more about Tropheus sp. “Crescentic” and Brichard’s Tropheus here.

Fig. 13. Tropheus sp. “Crescentic” at Segunga (northern shore), depth 5 metres. This particular geographical variant ("Yellow stain") was discovered by Horst Walter Dieckhoff in 1987 and labelled as originating from Isonga (Konings 1988: 124, fig. left, third from top).
Fig. 14. Tropheus sp. "Crescentic" at the northern side of Kibwe Bay. Depth 3 metres. The northern geographical variants of the species, from south of Malagarasi River to Maswa, exhibit some degree of a yellow vertical band behind the head. These variants were discovered by Wolfgang Staeck in 1974.

Suggestion on how to cite this blog article;

Karlsson, M. and Karlsson, M. (2015) A glimpse of the complexity of the genus Tropheus. African Diving Blog. Available from: http://blog.africandivingltd.com/2015/05/a-glimpse-of-complexity-of-genus.html (accessed [day] [month] [year])


African Diving Ltd (1990) Tropheus spec ”golden kushangaza”. Ciklidbladet, 23 (7): 1 (cover). Nordiska Ciklidsällskapet.

Evers, H.-G. (2015) Eine Prinzessin in Gold. Amazonas, 11 (2): 8-9; nr. 58. Natur und Tier-Verlag, Münster. Available from: http://www.amazonas-magazin.de/Produktinformation.790.0.html?&tt_products=4370 (accessed 18 March 2015)

Jonas, B. (ed.) (2015) Lake Tanganyika Conservation - Conservation of rare or endangered species of Lake Tanganyika. Available from: http://www.lake-tanganyika-conservation.com/ (accessed 18 March 2015)

Karlsson, M. and Karlsson, M. (2012) Tropheus moorii “Murago”. Ciklidbladet, 45 (2): 6-15. Nordiska Ciklidsällskapet.

Karlsson, M. and Karlsson, M. (2014a) Tropheus moorii “Murago” and its equal from Tanzania. African Diving Blog. Available from: http://blog.africandivingltd.com/2014/05/tropheus-moorii-and-its-equal-from.html (accessed 17 March 2015)

Karlsson, M. and Karlsson, M. (2014b) Lake Tanganyika Cichlids in the Wild: Tropheus moorii “Murago Tanzania” [video]. Production MMK Sweden. https://www.youtube.com/watch?v=DA4MCypRH4Y (accessed 17 March 2015)

Karlsson, M. and Lundblad, J. (1990) En hälsning från Tanganyikasjön. Ciklidbladet, 23 (6): 24-30. Nordiska Ciklidsällskapet.

Konings, A. (2014) The demise of a Tanganyika cichlid. The Cichlid Room Companion. Available from: http://www.cichlidae.com/section.php?id=279 (accessed 17 March 2015)

Leoni, L. (2015) Acquariofili. Santi o peccatori? (Prima parte). Mahengechromis - Divagazioni di un ciclidofilo. Available from: http://mahengechromis.blogspot.it/2014/12/acquariofili-santi-o-peccatori-prima.html (accessed 17 March 2015)

Mallet, J. (2005) Hybridization as an invasion of the genome. Trends in Ecology and Evolution, 20 (5): 229-237.

Salzburger, W., Baric, S. and Sturmbauer, C. (2002) Speciation via introgressive hybridization in East African cichlids? Molecular Ecology, 11 (3): 619–625.

Seehausen, O. (2004) Hybridization and adaptive radiation. Trends in Ecology and Evolution, 19 (4): 198–207.

Staeck, W. (2015) Cichliden von A bis Z - Tropheus sp. “Murago Tanzania”. DCG-Informationen, 46 (4): 80-84. Deutsche Cichliden-Gesellschaft e. V.

Ward, P. (2004) The father of all mass extinctions. Conservation in Practice, 5 (3): 12–17.

Zadenius, M. (1991) Importnytt - Nyheter från Åldes Akvarium. Ciklidbladet, 24 (9): 42-45. Nordiska Ciklidsällskapet.

Zadenius, M. (1991) Importnytt - Nyheter från Åldes Akvarium. Ciklidbladet, 24 (8): 36-38. Nordiska Ciklidsällskapet.

Zadenius, M. (1992) Tanganyikasjöns skönheter - Tropheus-släktet, del 2. Ciklidbladet, 25 (1): 14-22. Nordiska Ciklidsällskapet.

Zadenius, M. (1995) Importnytt från Åleds Akvarium AB. Ciklidbladet, 28 (1): 42-43. Nordiska Ciklidsällskapet.

Zadenius, M. (1997) Importnytt Trollenäs Zoologiska. Ciklidbladet, 30 (3): 48-49. Nordiska Ciklidsällskapet.

For additional references, see the literature list in the blog article about Brichard’s Tropheushttp://blog.africandivingltd.com/…/kushangaza-at-halembe-in…