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Review of Lake Tanganyika Cichlids in their Natural Habitat (3rd edition)

By , 2015. printer
Published
Mark Smith, 2008

Classification: Publications.

Tanganyika Cichlids in their Natural Habitat (3er edition) Tanganyika Cichlids in their Natural Habitat (3er edition).

The long awaited 3rd edition to Ad Konings’ Lake Tanganyika Cichlids, published in March 2015, has been a highly anticipated book for many Lake Tanganyikan cichlid aficionados. It has been over 17 years since the 2nd edition was published in late 1998, and before that, the first edition in 1988. These three books have ridden the wave of interest and ever evolving understanding and exportation of these cichlids, and this latest edition certainly maintains that momentum. Before beginning my assessment of this monumental 3rd edition, I would like to preface what is to follow by saying that I have greatly valued Konings’ contributions on the cichlids from Lake Tanganyika and Lake Malawi, as well as many other works he has published. A lot of time and research has gone into his writings, coupled with extensive underwater photography, all combined to come together into a unique and spectacular work for all of us hobbyists, and overall naturalists, to enjoy and learn from.

I would like to start out by saying what is noteworthy about this book. At 408 pages and over 1,200 photographs showcasing most of Lake Tanganyika’s cichlids, supplementing an extensive, yet not exhaustive, amount of research/information, this is the most comprehensive book ever published on the cichlids from this lake, and one to recommend to all hobbyists who have ever maintained Lake Tanganyikan cichlids, or is contemplating doing so. Konings has meticulously researched many current scientific papers on various aspects of Lake Tanganyikan cichlid origins, taxonomy, history, etc in order to have put together a rich textual work unlike no other. As a result, there are a lot of new ideas presented, which is what Konings is well known for. Some species names have been revalidated from synonymy in the past, while other recently described names have either been confirmed, or rejected.

Each and every book this author has published through Cichlid Press has been a true work of art, with spectacular photographic standards we have come to enjoy. So, when this new book failed to continue that line of high quality photography, due to the method of how the book was published, in a print-on-demand format in matte style, I can say that the quality of the images throughout this book are less than stellar. Having the photographs of this edition published in a matte finish and not in glossy format as in his previous books, has resulted in photographs that look somewhat hazy and less crisp. It is a huge departure from what we are all used to seeing in this author’s previous books. There is, of course, a reason for that, which has to do with the high cost of making quality books these days. Most hobbyists apparently do not purchase and build up much of a personal library, something that is quite unfortunate for those who profess a rich love of fishes. Excuses abound and become nauseating after a time. This, in turn, has caused authors like Konings to be more cautious in wanting to invest a lot of their money into thousands of printed copies that will not sell. In addition to that, Konings has indicated that he has no more space at his residence to hold onto several thousand copies of any new book he may get published. The end result is either this style of book, or nothing at all. At this point, I will take this book over nothing at all, especially for what the textual aspect has to offer.

I do have my own opinions on various species identifications that I, as well as many other hobbyists seriously devoted to Lake Tanganyikan cichlids around the world, as well as ichthyologists specializing in Lake Tanganyikan cichlids, do not agree with. An initial starting point is that there are several validly described species of cichlids from Lake Tanganyika that are no longer recognized as such in this book, not dismissed from officially published works critical of various species descriptions, but primarily from Konings own opinions. I tend to be a bit more cautious when it comes to invalidating a validly described species. That being said, there have been some descriptions that tempt one to perceive that they are the result of an extreme form of “splitting” or just plain poor research and analysis, such as for Cyphotilapia gibberosa, Neolamprologus cancellatus, Tropheus polli and Lepidiolamprologus nkambae.

" This is the most comprehensive book ever published on the cichlids from this lake "

I don’t relish this next section of this review, but it is necessary due to substantial changes/alterations on species identifications from the author’s perspective, as well as missing information on various species, not including certain undescribed species, and incorrect or outdated information that I take issues with, and for which will begin to cause confusion in the hobby if not challenged. Before jumping into these matters, the back cover states that the author has made over 750 hours of underwater observations. This same number of hours of underwater observations were stated in the 1998 2nd edition back cover, and that number remains the same in this 3rd edition. Surely that number is substantially greater now. In addition to that, the cover photograph is sort of a tease, as it is the only glossy image of a cichlid in the entire book.

I will give several examples of such changes, and omissions, which I bring to the attention of the reader, which are by no means exhaustive. One of my all time favorite species from the lake is Baileychromis centropomoides. Konings, on page 386, stated that the ichthyologists, Bailey and Stewart, who described this fascinating species, as well as from Adrian Indermaur, indicated that it is only found at two locations, namely Hore Bay and Sumbu Bay in the Zambian section of the lake. In my opinion, this is not entirely accurate. Bailey and Stewart never mentioned these two locations in their description of this species, and I can only assume that this is where Dr. Indermaur found them. Bailey and Stewart on August 5, 1977 published their description of this species in Occasional Papers Of The Museum Of Zoology, University of Michigan, number 679. In it, the authors stated that this species was collected just north northwest of Musende Bay, about 3 to 4 Kilometers west of Mpulungu, near Nsumbu, near Mpulungu, at Mwela, at Chituta Bay; and at Musende Bay, with a total of 13 specimens being collected from September 1970 to November 1972 by Kendall, Bailey and Ellis. Also, Konings informed me personally that the Karlsson brothers caught this species somewhere along the Tanzanian shoreline several years ago. This species’ natural distribution may encompass a large portion of the lake wherever suitable habitat is present.

Regarding Trematochromis benthicola, I had a paper published in early December 2014 in Tanganika Magazyn # 15, pages 62 through 73, specifically describing their spawning behaviour on pages 70 to 71. However, on page 215, last paragraph left column, Konings states that spawning has not yet been observed. I understand that he may have finished the text for this book before early December, but he was aware of myself having worked with this species for several years and was aware that I had spawned it repeatedly. He also states that males may reach 25 cm. My males, after nearly 5 years of age, never grew beyond 17 cm total length.

I don't see Julidochromis marksmithi as a variant of J. regani as this author does. Its pattern and behavior are clearly distinct from its closest relative, J. regani. At the fringes of its natural range, it may have hybridized with allopatrically occurring J. regani forms, thereby obscuring the uniqueness of J. marksmithi at the extreme northern and southern areas of its distribution. It may thus seem like a gradual transition from J. regani to J. marksmithi at these northern and southern limits. It is this transition of color forms which Konings has mentioned as justification for saying that the species recently described as J. marksmithi is nothing more than a color variant of J. regani.

Haplotaxodon trifasciatus is a species not acknowledged as being valid by the author, and is viewed as a synonym of H. microlepis. Haplotaxodon trifasciatus was described in 1999 in Copeia, #1, in a paper entitled New Species of Haplotaxodon (Perciformes: Cichlidae) from Lake Tanganyika, Africa by Tetsumi Takahashi and Kazuhiro Nakaya. Admittedly, their paper was not as detailed as it could have been, but nevertheless it was recognized as different from H. microlepis in a study by Muschick, Indermaur, and Salzburger, (2012). This is a classic case of cryptic speciation, and when compared side by side as juveniles, both H. trifasciatus and H. microlepis are nearly impossible to tell apart, except for the patterning in their unpaired fins and perhaps some subtle sheen-like patterning difference along the body. (This is reminiscent of very small juvenile Altolamprologus compressieps and A. calvus, which are impossible to tell apart if they are less than 15 mm total length). The real difference comes into play once they reach their adult size, with H. microlepis attaining approximately 35 cm total length while H. trifasciatus attaining approximately 23 cm total length. H. microlepis has a deeper body, somewhat more protruding snout, and tiny iridescent light blue flecks in the unpaired fins and over the entire body. In H. trifasciatus, it has a deeper head, more blunt snout, larger eye in relation to the size of the head, and a lightly colored irregular-ish horizontal bar along the middle of the dorsal fin. The body also has the subtle purple and yellowish horizontal barring along the entire length of the body. There is also a putative third species, with a much deeper body form than either of the two aforementioned species, which that was not mentioned in this edition.

Simochromis pleurospilus is viewed as a variant of S. babaulti, primarily due to a novel way to apparently determine what constitutes a unique species, which is based upon gill flukes, as reported by Van Steenberge (2014). Van Steenberge has posited that gill flukes are very host specific, so much so, that a particular species will only inhabit a particular cichlid species. And so, as the assumption goes, if two somewhat different looking cichlids possess the same gill fluke species, then they must belong to the same cichlid species. On the other hand, if two similarly looking cichlid species possess different gill fluke species, then it indicates they are two distinct cichlid species. That information is so new, that I would like to personally see additional specialists chime in on host specificity for such parasites. Does the same parasite parasitize closely related, yet distinct species, or are they so exacting in their host requirements that they will only invade a unique species? The all encompassing behavior of cichlids, and how these parasites are passed onto their progeny would be interesting to ascertain. Also, Van Steenberge may have indicated a reduced gill fluke host specificity with deep-water species from the lake. On another related yet different note, perhaps also recording the sounds that these cichlids make would also be another tool in deciphering their relatedness? Three photos of S. pleurospilus are depicted on page 232 which are those that display red dots on their flanks. This species is distinct from Simochromis babaulti in my opinion. Assuming that being the case, the implication suggests that specific gill fluke species can occupy at least two differing, yet closely related, cichlid species.

“Lamprologus” meleagris is considered a synonym of L. stappersi. This synonomy has also been presented in Konings 1998 2nd edition. I am not convinced. His argument seems like it makes sense, but until one goes to the exact river from where L. stappersi was found, or even directly in front of the river mouth and collect the cichlids from these locations, his reasoning is just a well thought out assumption that still has not been proven. I prefer to wait until such time as Konings, dedicated hobbyists, or ichthyologists go to the Sambala River and collect in its lower reaches and just out in front of it, and record what is collected.

On page 149, bottom left paragraph, Konings states that Lepidiolamprologus elongatus exhibits no geographical variation, which I interpret to mean no geographical color variation. This is hard to understand, that after 25 years, with different color variants coming out of the lake, that he would make such a statement. In addition to that, the Karlsson brothers Lakeside Articles magazine (Karlsson et al., 2013), showcase at least 6 different color variants of this species on pages 3, 12, 14, 15, 19, and the back cover. Some have a yellow cast to the body, while others vary substantially in the number and size of the white spots along the length of the body. Some variants have over twice the number of body spots than other variants.

Some of the above species, particularly C. gibberosa, N. cancellatus, L. nkambae and T. polli, were all poorly described, thereby adding to the confusion, and I think Konings has done a great job in explaining why these formally described species are suspect, if not outright invalid. Let me just say that, as is now apparent with the cichlids of Lake Tanganyika hybridizing for millennia and even today, it behoves those who describe the cichlids from this lake to be aware of this fact, and to tread cautiously, and at the least, to provide compelling and detailed reasoning behind each formal description. We look for ways to understand what is going on, and I applaud Konings for making persuasive, compelling arguments for his positions, even though I find some hard to swallow. One suggestion argues that a particular species may be a dwarf version of a particular, known species (see page 167 and 169 for what Konings calls a dwarf version of Julidochromis marlieri, but more commonly known as Julidochromis sp. “Gombi Transcriptus”, from Kombe, Zambia). Another argument will indicate that a particular species is a deep bodied version of a known species (see page 171 for what Konings calls a deep bodied version of Perissodus microlepis, which in my opinion is likely an undescribed species). These two examples appear to be too simplistic an explanation for what may be going on, and I get the impression that such reasoning is given to support a particular presupposition, or is presented just to have something to say about a given species that is hard to fully comprehend in terms of what it truly may represent. It seems that much more is going on, a lot of which we simply still do not understand, and to fit what is seen into neat and easy to understand formulas, misses the unique, subtle differences between species from this lake, a lake famous for such nuanced/subtly differing species, that frequently live side by side, such as is seen in Neolamprologus furcifer and Neolamprologus timidus.

Neolamprologus sp. 'ventralis kasanga' when it was first exported by Toby Veal, in 2002, was then also seen as an undescribed species by Konings. It was initially exported under the name Neolamprologus sp. 'red dorsal ventralis'. It apparently lives at the same depth range and habitat as does N. ventralis. This species was eventually referred to by Konings as N. ventralis in Back To Nature Guide to Tanganyika Cichlids 2nd Edition, published in 2005. There are substantial differences between Neolamprologus sp. 'ventralis kasanga' and N. ventralis. In N. sp. 'ventralis kasanga' males and females possess much shorter ventral fins, the female is much smaller and less colorful than the male, (whereas in N. ventralis both male and female are the same size and coloration at some locations, while at other locations the male may be slightly larger and somewhat more colorful), their dorsal fin and anal fin height is shorter, their body is deeper, snout shorter, and fewer scales in a lateral line 31 to 32 for N. sp. 'ventralis kasanga' vs. 34 to 36 for N. ventralis. Lastly, N. 'ventralis kasanga' is a much milder mannered Lamprologine than N. ventralis is in captivity, which made it very easy to spawn in captivity, than for N. ventralis. In addition to that, there is a vertically barred undescribed Neolamprologus species similar to N. ventralis, see page 180 middle photo, which is viewed as a variant of N. ventralis by Konings. This too, is likely an undescribed species.

Regarding the gold head variant of Neolamprologus cylindricus, the author makes an assumption that it may be a hybrid between N. leleupi and N. cylindricus. This seems like a preposterous thing to say. Even though the exact collecting locality is unknown (however, it is hinted at that it may hail from Isanga, Zambia in the second edition from 1998) it nevertheless is N. cylindricus, as is evidenced in the adult form, being identical to the variant found at Nkondwe Island, Tanzania. The gold head N. cylindricus, is given this name due to the juveniles possessing orange blotches over the entire head, nape and front third of the body, much like that seen in juvenile Neolamprologus sp. 'falcicula cygnus'. In both species, the orange blotches fade completely away as they mature into adults.

Another species that stood out to me as likely not being correctly identified is what the author is calling Neolamprologus mustax from Kiku Congo, on page 136 top photo, showing a very large eyed, very blunt snouted, possibly undescribed species.

Lepidiolamprologus kamambae was given subspecific status to L. kendalli by the author, following Tawil et al., 2013. In my opinion it should revert back to L. kamambae.

This one is truly baffling. On page 220, upper photo, is what is purporting to be a Lepidiolamprologus pleuromaculatus from Kiriza, Congo. Not only is the specimen in the photo twice as slender as L. pleuromaculatus from the Burundian coastline, the patterning in the unpaired fins and snout shape differ markedly from L. pleuromaculatus. In fact, it is even more slender than L. attenuatus! This one should in my opinion best be labeled as Lepidiolamprologus cf. attenuatus from Kiriza, Congo for the time being.

Neolamprologus crassus now N. pulcher? This may be a situation of active hybridization or recent introgressive hybridization events, but to call N. crassus a variant of N. pulcher at this stage of the “evolutionary” process stretches all credulity. It is not even agreed upon that N. brichardi is a synonym of N. pulcher. The authors of the paper (Duftner et al., 2007) that propose N. brichardi to be synonymous with N. pulcher even leave some doubts that these two species are one and the same, although the evidence they give strongly points to them being the same species. Admittedly, this is a mysterious and difficult one to fully understand and I get the impression that the last word on these species has not yet been written.

For the genus Trematocara, the photo on page 375, bottom photo, purports to show a Trematocara variabile, but in my opinion it is a female T. marginatus. Also, an intriguing image of a completely new Trematocara species that has not made its way into this new edition, was showcased on the Cichlid Room Companion forum in a photo probably taken by Benoit Jonas back in January 2007, stated to had been taken while visiting the Congo side of the lake at Makobola. It shows a Trematocara species with incredibly large eyes. It also has interesting golden flecking on the lower half of the body, and an orange/yellow anal fin. A clear stripe runs horizontally through the back half of the dorsal fin.

The Petrochromis polyodon complex. In my opinion it would have been better to say P. cf. polyodon for the multitude of specimens found throughout the lake (see pages 54 through 59), rather than saying that they are all P. polyodon. None of these races have been collected and studied morphometrically, molecularly, or even for their gill flukes, if in fact is it as reliable as Van Steenberge suggests. In the text, the author is not completely sure of them being all the same species. In order to be a bit more cautious on labeling less than clearly defined species, the liberal use of the “cf.” designation would have been more useful for this group of cichlids, as well as for many species depicted in this edition, since the author frequently indicates doubt as to the veracity of various species, but then goes on to confidently say they are “thus and so” species.

There are many other formally described species that the author has decided are not valid and I just list them here without going into further detail on them: Neolamprologus chitamwebwai is considered a synonym of Neolamprologus falcicula; Telmatochromis brachygnathus is considered a polymorphic variant/synonym of Telmatochromis temporalis; Cyphotilapia gibberosa is considered a synonym of Cyphotilapia frontosa; Neolamprologus cancellatus is considered a likely hybrid; Lepidiolamprologus nkambae is considered a synonym of Lepidiolamprologus kendalli; Neolamprologus helianthus is considered a synonym of Neolamprologus splendens; Neolamprologus brichardi is considered a synonym of Neolamprologus pulcher; Tropheus polli is considered a synonym of Tropheus annectens: and Neolamprologus petricola is now considered a synonym of Neolamprologus modestus. Some of these synonymised species make a lot of sense, while others are hard to accept at this point.

Beyond that, it seems to me that most other species are dealt with in an accurate, innovated manner. However, other Lake Tanganyika experts, such as Patrick Tawil, have coined names for undescribed species, for which this author has changed, an example being for an undescribed Cyprichromis species, namely C. sp. 'dwarf jumbo' that Tawil proposed back in 2008. In this 3rd edition, the author renames it Cyprichromis sp. 'leptosoma kigoma’. This will only serve to cause unnecessary confusion, and this undescribed species should continue to be referred to as C. sp. “dwarf jumbo” in my opinion.

Perissodus multidentatus Perissodus multidentatus male in the aquarium. Photo by Mark Smith. (02-Jun-2004).

In some instances, photographs of higher quality specimens should have been used, such as for Perissodus multidentatus. There is somewhere an aquarium photo of a freshly caught specimen with virtually no prominent coloration or elaborate finnage that this species is famous for. My photo of a fully colored and physically developed P. multidentatus on The Cichlid Room Companion species profile would have been a better choice to show what this spectacular species looks like. Also, what is being called “Lamprologus” ornatipinnis from Kigoma, Tanzania, photographed by Dieckhoff, on page 342, bottom photo, shows two specimens with fins folded down, not showing the most prominent characteristic that this, in my opinion, undescribed species from Kigoma is also famous for. Again, my images, posted on The Cichlid Room Companion catalog, of a male and female, show this exact species from Kigoma with unpaired fins fully extended, showcasing their beautiful black and white striping in the dorsal and anal fins. In addition, I find it unlikely that this is the real L. ornatipinnis. The holotype of this species was collected one kilometer south of Mtoto, on the west coast of the lake, and the largest specimen recorded was 78 mm total length. The maximum size of the Kigoma specimens I kept was significantly smaller at approximately 62 mm for the male, and 52 mm for the female.

In other instances, various species were entirely omitted from this 3rd edition. There was a very pretty Lepidiolamprologus cf. meeli imported over 25 years ago without a specific location given other than that of Congo. It possesses attractive speckling on the body and black and white markings on the fins, easily making it the prettiest species in the L. meeli guild to date. Unfortunately, it was not included in this new edition, or in any other book on Lake Tanganyikan cichlids that this author has ever published. A picture of this species can be viewed in my book, Lake Tanganyika Cichlids, 2nd edition published in 2007, on p. 58, lower right photo. Also, on page 91 of my book, a photo by Thomas Anderson shows an undescribed cichlid collected from deep water in Chituta Bay, Zambia, as a by-catch of Xenotilapia nasus. It likely represents a new genus of cichlid, most likely from the tribe Ectodini, and this was not included in this 3rd edition, even though the author was aware of this species for many years. No image of a blotched N. leleupi that Rene Kruter photographed many years ago either (Krüter, 2012:7, lower right photo).

So, is this book worth the $85 dollar price tag? I would say yes with reservations. Just be prepared to be met with photographs that are not what you have come to expect from Cichlid Press. As I look through this new edition more and more, the photographs are beginning to grow on me, but will never have the same punch that glossy images give off. I just hope this author does not attempt to come out with a 5th edition to his Lake Malawi cichlids book in a similar manner. In terms of the multitude of, in my opinion, incorrectly identified species, well, I leave that up to you to chew your way through and make sense of. And, as for those species that never made it into this book, hopefully someday we will be able to see them all together in a future edition. On a side note, it would have been nice to have seen more images of non-cichlids and aquatic invertebrates. They seldom receive decent coverage that it would nice for a change to really attempt to cover their diversity with a comprehensive selection of images and information

References (15):

Citation

Smith, Mark. (March 30, 2015). "Review of Lake Tanganyika Cichlids in their Natural Habitat (3rd edition)". Cichlid Room Companion. Retrieved on December 17, 2018, from: https://www.cichlidae.com/article.php?id=442.