ACTA ICHTHYOLOGICA ET PISCATORIA (2011) 41 (3): 201–214 DOI: 10.3750/AIP2011.41.3.08

A RECENT INVENTORY OF THE OF THE NORTH-WESTERN AND CENTRAL WESTERN COAST OF (DEMOCRATIC REPUBLIC CONGO)

Maarten VAN STEENBERGE 1,2*# , Maarten P.M. VANHOVE 1,2# , Donatien MUZUMANI RISASI 3, Théophile MULIMBWA N’SIBULA 3, Fidel MUTEREZI BUKINGA 3, Antoine PARISELLE 4, Céline GILLARDIN 2, Emmanuel VREVEN 1, Joost A.M. RAEYMAEKERS 2, Tine HUYSE 2, Filip A.M. VOLCKAERT 2, Venant NSHOMBO MUDERHWA 3, and Jos SNOEKS 1,2

1 Ichthyology Unit, African Zoology Department, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080, Tervuren, Belgium 2 Laboratory of Diversity and Systematics, Department of Biology, Katholieke Universiteit Leuven, Charles Deberiotstraat 32, B-3000, Leuven, Belgium 3 Section de Zoologie, Département de Biologie, Centre de Recherche en Hydrobiologie, B.P. 73, Uvira, Democratic Republic Congo. Via B.P. 254 Bujumbura, Burundi 4 ISE-M, UMR5554 CNRS, UR226 IRD, Université Montpellier II – CC 063, F-34095, Montpellier Cedex 5, France

Van Steenberge M., Vanhove M.P.M., Muzumani Risasi D., Mulimbwa N’sibula T., Muterezi Bukinga F., Pariselle A., Gillardin C., Vreven E., Raeymaekers J.A.M., Huyse T., Volckaert F.A.M., Nshombo Muderhwa V., Snoeks J. 2011. A recent inventory of the fishes of the north-western and central west - ern coast of Lake Tanganyika (Democratic Republic Congo). Acta Ichthyol. Piscat. 41 (3): 201–214. Background. Despite the importance of Lake Tanganyika’s biodiversity for science and the livelihoods of the riparian people, high-resolution surveys of the biodiversity are sparse and fragmentary, especially along the western (Congolese) shoreline. The coast suffers locally from intensive human activities and lacks adequate pro - tective measures or nature reserves. However, in view of the intra-lacustrine endemism of this fish fauna, con - servation needs to be managed lake-wide at a fine scale, necessitating detailed inventories on fish species distri - bution. The study aims at updating knowledge on fish diversity and distribution along the north-western and cen - tral western shores of Lake Tanganyika. Materials and methods. Fish specimens were collected using gill- and seine nets, by snorkelling and SCUBA diving, and through purchases on the local markets. Results. Over 28 locations were sampled, and 84 - and 30 non-cichlid fish species (belonging to Protopteridae, Clupeidae, Cyprinidae, Alestidae, Claroteidae, Clariidae, Malapteruridae, Mochokidae, Poeciliidae, Latidae, and Mastacembelidae) collected. Conclusion. Our records substantially expand the known range of fish species in a range of habitats. As numer - ous specimens are hard to assign to nominal species, a taxonomic revision of a number of genera is underway. It should take into account intraspecific geographic variation. Keywords: Africa, ancient lake, biodiversity, Cichlidae, ichthyofauna, Mastacembelidae, Mochokidae, Teleostei

INTRODUCTION River and is therefore regarded as one of the sources of the Lake Tanganyika offers vital resources including Congo Basin. Lake Tanganyika is the deepest and the old - food, household water, and transportation to an estimated est of the African Great Lakes (Cohen et al. 1997). It is 10 million people in the riparian countries, with an eco - home to about 250 species of Cichlidae, the world’s mor - nomic impact reaching far inland (Mölsä et al. 1999). The phologically and genetically most diverse cichlid assem - Democratic Republic Congo (DRC) part of Lake blage, and 75 non-cichlid fish species (Snoeks 2000). The Tanganyika extends over 14 800 km² (45%) of the lake’s in the lake are polyphyletic and include ancestral surface and 795 km (43%) of its perimeter. The lake’s lineages (Nishida 1991, Salzburger et al. 2005, Koblmüller only outflow, the Lukuga River, feeds into the Congo et al. 2008). Its diversity makes Cichlidae in general

* Correspondence: Drs. Maarten Van Steenberge, Ichthyology Unit, African Zoology Department, Royal Museum for Central Africa, Leuvensesteenweg 13, B-3080, Tervuren, Belgium, phone: +32 (0)2 7695632, fax: +32 (0)2 7695642, e-mail: [email protected] . # Both authors contributed equally. 202 Van Steenberge et al. and those of Lake Tanganyika in particular an important key Lake Kariba to support a pelagic fishery (Bell-Cross and group for understanding evolution in lower vertebrates Bell-Cross 1971). For littoral fish communities, pollution, (Kornfield and Smith 2000, Koblmüller et al. 2008). Most mosquito net fishing and increased sedimentation (following taxonomic work on Lake Tanganyika cichlids was per - deforestation) pose serious threats (Cohen et al. 1993, Hori et formed by Boulenger (1915) and Poll (1956, 1986). al. 1993, Allison et al. unpublished ** , Donohue et al. 2003). Snoeks (2000) pointed out that since then, discovery and Adverse effects of increased turbidity on the cichlid fauna description of new varieties and species have mostly been have already been shown for amongst others Lake Victoria carried out by ornamental fish hobbyists. Most informa - (Seehausen et al. 1997, Galis and Metz 1998). Although the tion regarding their distribution is scattered through the stakeholders are well aware of problems, legislation is not aquarium literature (e.g., Brichard 1989, Konings 1998, up-to-date (Fermon 2007) and not any nature reserve has Schupke 2003). Lake Tanganyika also excels in endemic - been designated in the Congolese part of the lake (Hori et ity and radiation of non-cichlid fishes (Snoeks 2000, al. 1993), while the regions around Uvira and Moba are Salzburger et al. 2002). Indeed, the lake is home to subject to intensive human activities (Mölsä et al. 1999). species flocks belonging to several other fish families, As many fish species have limited distributions within like Latidae (Coulter 1976), Mastacembelidae (Vreven Lake Tanganyika and as species communities are highly and Snoeks 2009, Brown et al. 2010), and Mochokidae structured on a small geographical scale (Sturmbauer 2008), (Day and Wilkinson 2006, Koblmüller et al. 2006). it is important that conservation efforts are well targeted and Moreover, several invertebrate taxa radiated within Lake spread across the lake. Unfortunately, few data are available Tanganyika, such as atyid prawns (Fryer 2006), platytel - on species distributions at a high resolution (Kawabata and phusid crabs (Marijnissen et al. 2006), thiarid (West and Mihigo 1982, Hori et al. 1983, Fermon 2007), and often they Michel 2000, Michel et al. 2004) and thallasoid (Wilson et are only found in the grey literature. In 1992, 1995, 2001, al. 2004) gastropods, copepods (Coulter 1991a), ostracods and 2008, the Royal Museum for Central Africa (RMCA) (Martens and Schön 1999, Wouters and Martens 2001) and the Royal Belgian Institute for Natural Sciences, and possibly Eunapius (spongillid Porifera) (Erpenbeck together with local and international partners, joined et al. 2011). forces in expeditions exploring most of the Tanzanian and Dominating the pelagic zone and serving as the main Zambian coast, and resulting in large RMCA collections target for fisheries are the clupeids Limnothrissa miodon * on the lake’s ichthyofauna. Political instability in the and Stolothrissa tanganicae Regan, 1917 and their latid region has hampered sampling and hence obstructed sci - predators Lates stappersii and economically less impor - entific data gathering especially from the DRC (e.g., tant L. mariae ; L. angustifrons ; and L. microlepis (see Schupke 2003 for Tropheus Boulenger, 1898). The aim Coulter 1991b). Cichlids, however, dominate in the littoral here is therefore to provide a formal update zone, certainly in rocky habitats (Brichard 1989). of our knowledge on the diversity and distribution of fish - Substrate has a considerable influence on the species com - es over a variety of habitats along the north-western and position of shore communities, with the rocky littoral central-western Tanganyika coast. being characterised by a higher degree of endemism (Brichard 1978, Konings 1998). Although ecological net - MATERIALS AND METHODS works of species interactions render those communities The 28 sampling sites (Table 1, Fig. 1) were mainly rather stable in spite of some local perturbation, they are located in the rocky littoral, although specimens were also not adapted to “new” disturbance like pollution or invasive collected in intermediate habitats, sandy beaches, adjacent species. Hence, conservation efforts are important (Hori et swamps, and the Lukuga outflow of the lake. Fish were al. 1993). Whereas the high fishing pressure on the pelag - caught while snorkelling and diving, using gill nets of ic (Mulimbwa and Shirakihara 1994, Mölsä et al. 1999, mesh sizes 8, 10, 12, and 15 mm on rocky shores, and Mulimbwa 2006) and littoral (Allison et al. unpub - with seine nets at sandy beaches. At Bulumba Island and lished**) fish stocks might jeopardise sustainability, cli - Mtotokainda, gill nets were placed overnight and at mate change represents another threat to pelagic fisheries Luhanga and Bemba, specimens from the deeper littoral (O’Reilly et al. 2003). The introduction of exotic species zone were collected by SCUBA diving (Table 1). should be considered and avoided (Hori et al. 1993). Additional specimens were obtained from fishermen and However, successful establishment of exotics appears to fish markets. Specimens were identified to species level remain limited in Lake Tanganyika so far; niches are well in situ and in the laboratory, following Poll (1978, 1986), occupied and hence buffered (Hall and Mills 2000). Yamaoka (1983), Brichard (1989), Eccles (1992), Vreven Moreover, the physicochemical differences between the (unpublished *** ), Wright and Page (2006), and Retzer lake and surrounding water bodies (Coulter 1991a) might (2010) and by comparison with type material. Fishes were be a factor hampering spontaneous immigration. The lake collected from the Congolese lakeshore under mission has served as a “source” for the introduction of two of its statement (ordre de mission) no. 013/MNRST/CRH- endemic species, the freshwater sardine Limnothrissa U/2010 from the Ministère de la Recherche Scientifique et miodon and the killifish Lamprichthys tanganicanus , into Technologique – Centre de Recherche en Hydrobiologie Lake Kivu (Nshombo and Lushombo 2010). Limnothrissa (CRH, Uvira). A reference collection has been stored at miodon was also introduced into artificial lakes such as the CRH (Uvira, DRC) and at the RMCA (Tervuren,

* Authorities of the species examined in the presently reported study are provided in Tables 2 and 3 . ** Allison E.H., Paley R.G.T., Ntakimazi G., Cowan V.J., West K. 2000. Biodiversity assessment and conservation in Lake Tanganyika. BIOSS final technical report. *** Vreven E. 2001. A systematic revision of the African spiny-eels (Mastacembelidae; Synbranchiformes). PhD thesis. K.U. Leuven, Leuven. Fishes from the Congolese coast of Lake Tanganyika 203

Belgium). Taxon and author names in this study follow tions, were seldom encountered. Cyprichromis microlepi - Seegers (2008) for claroteid catfishes belonging to dotus , a species that is mostly found below 10 m of depth, Bathybagrus Bailey et Stewart, 1984, was also only collected twice. At both occasions, this was and Eschmeyer and Fricke (2011) for all other fish species. at the leeward side of an island and far from the shore. This corresponds to Brichard (1989) who claimed that RESULTS Cyprichromis only occurs in very quiet water. In total we collected 84 cichlid- and 30 non-cichlid Lepidiolamprologus mimicus , an aggressive mimic species. Tables 2 and 3 provide an overview of species of C. microlepidotus (see Schelly et al. 2007) was collect - presence at the various localities. ed only together with its model. Moreover, the maximal This list is not intended to be a complete species inven - number of cichlid species collected (36 at Mukamba, tory of the collection sites. As the vast majority of speci - where the collection effort was the highest) is lower than mens were collected using gill nets that were placed close the number of species encountered on a 400 m² quadrant to shore, species that are known to occur in deeper waters, by Hori et al. (1983) (38) at Luhanga in the north-western and shell or cave dwellers are missing or underrepresent - and by Sturmbauer et al. (2008) (41 and 46) and Takeuchi ed. For example, furcifer , a common et al. (2010) (54) in the southern part of the lake. Raiamas cave dweller with a secretive lifestyle (Konings 1998) was moorii was only caught in turbulent water in intermediate only caught once. As it was only collected when nets were habitats, where visibility was low. The fisheries target left overnight, this agrees with Brichard (1989) who sug - species belonging to the Boulengerochromini, gested N. furcifer to possibly be a nocturnal species. Also Tylochromini, Tilapiini, Bathybatini as well as Lates the pelagic species belonging to Clupeidae, Latidae, and stappersii and Malapterurus tanganyikaensis were only Bathybates that are known to have lake-wide distribu - obtained through fishermen operating from sediment Table 1 Sampling localities in Lake Tanganyika with habitat types and collection method Sampling site (or locality) Date Coordinates Habitat type Collection method Nyangara wetland, 1 Rusizi floodplain 20 Mar 2010 3°20′S, 29°09′E (market) swamp from Kilomoni market 2 Luhanga 27 Mar 2010 3°31′04′′S, 29°08′57′′E rocky shore snorkelling, SCUBA 3 Bemba rocky shore snorkelling, SCUBA 26 Mar 2010 3°37′22′′S, 29°08′56′′E sandy beach 4 Munene with vegetation caught by hand includes overnight gill 5 Bulumba Island 25 Mar 2010 3°46′11′′S, 29°07′10′′E rocky shore netting 6 Lubumba 24 Mar 2010 3°58′54′′S, 29°06′32′′E rocky shore snorkelling 7 Cap Banza 22 Mar 2010 4°03′53′′S, 29°13′48′′E rocky shore snorkelling rocky shore 8 Kisokwe 23 Mar 2010 4°14′31′′S, 29°10′35′′E with pebbles snorkelling 9 Mukindu 25 Apr 2010 5°36′01′′S, 29°22′36′′E rocky shore snorkelling 10 Murega 24 Apr 2010 5°38′22′′S, 29°23′06′′E rocky shore snorkelling 11 Mtotokainda 25 Apr 2010 5°38′60′′S, 29°22′53′′E rocky shore overnight gill netting 12 Magogoro 23 Apr 2010 5°39′19′′S, 29°22′51′′E rocky shore snorkelling 13 Musinwa 25 Apr 2010 5°40′57′′S, 29°24′31′′E rocky shore snorkelling 14 Kalemie 23–24 Apr 2010 5°54′54′′ S, 29°11′39′′E Lukuga outfl ow purchased on market intermediate purchased from local 15 Mulembwe 9 Apr 2010 6°07′10′′S, 29°16′18′′E with vegetation fi shermen, overnight gill net 16 Cape Tembwe 10 Apr 2010 6°29′39′′S, 29°25′29′′E intermediate snorkelling 17 Kabulu 21 Apr 2010 6°39′32′′S, 29°29′36′′E intermediate snorkelling sandy beach purchased from local 18 Mpala 21 Apr 2010 6°44′53′′S, 29°31′59′′E with swamps fi sherman 19 Mugayo North 11 Apr 2010 6°46′42′′S, 29°33′30′′E rocky shore snorkelling 20 Mugayo 11 Apr 2010 6°46′51′′S, 29°33′42′′E beach seine net 21 Mukamba 14 and 16 Apr 2010 6°56′51′′S, 29°42′43′′E rocky shore snorkelling 22 Mtoto 15 Apr 2010 6°58′03′′S, 29°43′50′′E rocky shore snorkelling 23 Kapakwe 15 Apr 2010 6°58′27′′S, 29°44′05′′E rocky shore snorkelling purchased from local 24 Moba 13–20 Apr 2010 7°02′30′′S, 29°46′30′′E sandy beach fi shermen, seine net 25 Migenzi 16 Apr 2010 7°04′56′′S, 29°53′17′′E rocky shore snorkelling 26 Mufazi 13 Apr 2010 7°05′12′′S, 29°54′45′′E rocky shore snorkelling 27 Kyanza 19 Apr 2010 7°06′42′′S, 29°58′34′′E rocky shore snorkelling 28 Kikoti 20 Apr 2010 7°11′28′′S, 30°04′01′′E rocky shore snorkelling

F 204 Van Steenberge et al. shores. Several members of the ecologically diverse tribe S. dhonti (Boulenger, 1917) monotypic. After close exam - Ectodini (see Koblmüller et al. 2004) (e.g. , Ectodus ination of the type series, we also found an axillary pore descampsii , Grammatotria lemairii , Xenotilapia on the holotype and two of the six paratypes of S. irsacae . melanogenys , and X. sima ) as well as Mastacembelus Therefore, the status of this species should be reviewed. albomaculatus , were only obtained by seine netting at As the type specimen, and the only specimen, of S. dhon - sandy beaches. Species endemic to the Lake Tanganyika ti measures 395 mm total length (TL) and the largest spec - basin but not found in the lake proper were collected in imen available of S. irsacae measures only 161 mm (TL), the vicinity of the Lukuga outflow. These include Barbus comparing both species is difficult and additional materi - lufukiensis, Micralestes vittatus , Auchenoglanis tangani - al is required. canus *, and Haplochromis burtoni . From the central western coast, S. polli had hitherto not been collected, whereas S. petricola was only known DISCUSSION from Mtoto until now. Given that the two species are now The present study features among the first during the known from the extreme south, the extreme north and last decades to formally report on fish diversity and distri - from a few localities along the east coast, this suggests bution along the Congolese Lake Tanganyika coast. The that they have a lake-wide distribution. fish community composition differs substantially between Geographical variation versus species status in the sampling localities, pointing to different geographical paraphyletic cichlid genus . Recently ranges and habitat preferences (e.g. , regarding substrate) intraspecific geographic variation has increasingly been of the various taxa. Sampling strategy (e.g. , methodology, valued in cichlid taxa of the East-African Great Lakes sampling effort, weather conditions, time of the day) (Hanssens and Snoeks 2003, Risch and Snoeks 2008, influences which species are caught. Although the rocky Anseeuw et al 2011). In Lake Tanganyika, such variation littoral is clearly the most diverse Tanganyika habitat, is especially found in species from rocky shores, both in with the highest degree of endemism ( cfr. supra ), these colour pattern (Kohda et al. 1996, Konings 1998) and in results demonstrate the need to include as many habitat morphology (Risch and Snoeks 2008). As many Lake types as possible in biodiversity surveys of both cichlid Tanganyika species are described from a single locality, and non-cichlid radiations. Fine-scale insights in the assigning a geographical morph to a nominal species often species diversity of fishes and other biota across the vari - proves difficult. This was, for example, the case for some ous lake habitats can be used to propose additional areas for conservation, or to efficiently manage existing ones. In this regard, Sturmbauer (2008) proposed the establishment of many micro-scale protected areas that can be managed by local communities and that maximize the conservation of overall biodiversity. Additionally, practices other than protected reserves are recommended, such as coastal zone management (Allison et al. unpublished* *) and a ban on littoral fisheries (Mulimbwa 2006). Taxonomic problems and distribution in Synodontis mochokid catfishes. Some specimens belonging to the species-rich genus Synodontis Cuvier, 1816 were difficult to identify. The specimens referred to as S. aff. multipunctata 1 and S. aff. multipunctata 2 agree with the description of Wright and Page (2006), except that the occipito-nuchal shield is naked in these specimens whereas it should be covered with skin in S. multipuncta - ta . These specimens were classified as two species as they differ greatly in pigmentation pattern. The specimen iden - tified as S. aff. multipunctata 1 has many medium-sized spots on the dorsal but not on the ventral part of the body, whereas S. aff. multipunctata 2 has fewer, but larger spots that occur on the whole body. The specimen assigned to S. irsacae corresponds to the description of the species as provided by Wright and Page (2006), except for the axillary pores, which are clearly visible in this specimen. Matthes (1962) put S. irsacae into synonymy with S. dhonti , a suggestion which was later confirmed by Poll (1971). The absence of an axillary pore was one of the reasons leading Wright Fig. 1 . Map of Lake Tanganyika; numbers refer to sam - and Page (2006) to rehabilitate S. irsacae , rendering pled locations (Table 1)

* Editor’s comment: According to FishBase Auchenoglanis tanganicanus Boulenger, 1906 is a junior synonym of Auchenoglanis occidentalis (Valenciennes, 1840) The present authors, however, adhere to Eschmeyer and Fricke (2011) for nomenclature and more specifically for this species to Retzer (2010). ** See footnote on page 202. Fishes from the Congolese coast of Lake Tanganyika 205 of the Simochromis Boulenger, 1898 specimens collected. eretmodine Eretmodus Boulenger, 1898, two distinct Simochromis marginatus is a species described from five morphotypes were encountered along the western shore. specimens caught at Manga on the Ubwari peninsula (Fig. 1). South of Kalemie, the typical E. cyanostictus was collect - It was hitherto unknown from more southern locations ed, whereas to the north of the Lukuga River, a variant along the west coast. At the Kavala Islands (localities 11 with vertical bands across the flanks and a subterminal and 12), Simochromis specimens were assigned to S. mar - and wider mouth was found. The existence of two ginatus , following a detailed morphological analysis Eretmodus spp. was suggested by Verheyen et al. (1996) (unpublished data). The specimens did however differ and Rüber and Adams (2000). Rüber et al. (1999) looked from the description given by Poll (1956), as they lack the at mitochondrial sequences of all eretmodine species, and black band on the dorsal fin, a character used by Poll found six different lineages that did not correspond with (1956) to distinguish this species from its congeners. the nominal species. For E. cyanostictus , two lineages The best studied areas are situated at the northernmost were found: A and C, the distributions of which corre - and southernmost end of the lake’s shoreline. Therefore, spond with those of the two morphotypes encountered species formally described from populations collected at here. The undescribed northern species is listed here as either side could either be true sister species, or part E. cyanostictus “north”, following Konings (1998). of a continuum of geographical morphs. Konings (1998) It should, however, be mentioned that according to proposes the latter hypothesis for S. babaulti and S. pleu - Konings (1998), the “true” E. cyanostictus is to be found rospilus , the former being described from specimens from even further south and that E. cyanostictus from Uvira in the extreme North, the latter from Zambian spec - Kapampa, near Kiku (Fig. 1), has a colour pattern inter - imens in the extreme South. In Nelissen’s (1978) descrip - mediate between the northern and the southern form. tion of S. pleurospilus , geographic variation in S. babaulti A comparable situation was found in the lampro - was not included. Although he mentioned that both logines Poll, 1974 and species occur in sympatry along the southern coast, he did Boulenger, 1898. Both genera have a similar ecology and not compare S. pleurospilus with S. babaulti specimens rarely co-occur (Konings 1998). For Julidochromis , originating from the southern part for wanting of suffi - a specimen of J. marlieri was caught at Bemba whereas ciently sized southern S. babaulti specimens. As Nelissen further south, at Bulumba Island and Lubumba, J. regani (1978) did not mention any clear morphometric or meris - was found. While for the northern basin J. marlieri is tic characters that can be used to differentiate between known from Burundi and a few locations scattered along S. pleurospilus and S. babaulti , the difference between the Congolese side (Konings 1998, Allison et al. unpub - both species mostly lies in their colouration pattern: lished *), J. regani is found on the north-eastern S. pleurospilus has numerous small red spots arranged in (Burundese) side, off the Ubwari Peninsula and, as in this horizontal rows on the flanks, while only a few red spots survey, on some western localities to the north of it are present in S. babaulti . In specimens collected to the (Brichard 1978, Konings 1998). At localities further south of Moba, spots were always visible in the larger south, three distinct morphotypes of Chalinochromis were individuals, and these specimens were therefore identified collected. Currently, only two species have been as S. pleurospilus . In specimens collected to the north described within this genus, although some geographical of Kalemie, very few or no spots could be seen, which forms probably deserve species status (Konings 1998). justifies their assignment to S. babaulti . In specimens Chalinochromis brichardi and C. popelini can be distin - caught between Kalemie and Moba, spots were some - guished by (1) the shape of the caudal fin, which is round - times clearly visible and sometimes absent, even on large ed in the former and lyre-shaped in the latter, and (2) by individuals. Specimens from those populations were, until their colour pattern: C. popelini always has three com - further study, identified as S. babaulti . As colour patterns plete lateral bands, one at the base of the dorsal fin and are known to show a great deal of variation within sever - two over the lateral line and a dark spot on the base of the al Lake Tanganyika cichlid species (Konings 1998), one caudal fin, whereas C. brichardi has zero to two bands should avoid colour as the sole criterion for species delin - and lacks a spot on the caudal fin (Brichard 1989). The eation. From these examples, it is clear that the para - specimen collected at Musinwa off the Kavala Islands has phyletic genus Simochromis (see Koblmüller et al. 2010) a rounded caudal fin, lacks any bands on the flanks and is in need of revision. The same holds for the paraphylet - corresponds to the description of C. brichardi . All speci - ic genus Petrochromis Boulenger, 1898 and although mens collected between Kalemie and Moba have a lyre- specimens collected during this survey were identified shaped caudal fin and three bands on the flanks; they have according to the current , it should be noted that been identified as C. popelini . The two specimens collect - the genus’ species richness is drastically underestimated, ed further south at Kyanza and Kikoti have a rounded cau - with the nominal species P. polyodon alone containing dal fin and show only two clear bands on the flanks. several undescribed species (Konings 1998, Makasa and Moreover, they differ from C. popelini as they do not Snoeks 2003, Koblmüller et al. 2010). have a dark spot at the base of the caudal fin. Given that The need of taxonomic revision of Lake such specimens are known in the aquarium trade under Tanganyika’s cichlid fauna: examples from eretmo - the name C. sp. “bifrenatus” (Tawil 1986), these speci - dine and lamprologine cichlids. Of the monospecific mens are listed here as such. This record is interesting

* See footnote on page 202. 206 Van Steenberge et al. 2 8 e + + + 2 l b a 7 + + + + + T 2 6 + + + + 2 5 + + + 2 4 + + 2 3 + + 2 ) 2 + 1 2 1 0 1 2 + + + + + + + ( 2 e k 0 + + + + c 2 i r F 9 + + + d 1 n a 8 r + + + + 1 e y e 7 + + + + 1 m h c 6 s + + + 1 e E t i s g 5 n + + + g i 1 n i w l o p 4 l + + + + + + + + l 1 m o a f S 3 y + + + + 1 l i m 2 a + + 1 f y 1 b + + + + + + 1 d e i 0 f + + i 1 s s a l 9 + c , a k 8 + + i y n a 7 + + + g n a 6 + + T e k 5 a + + + + + + + + L n i 4 + , s e h 3 + + s i f d 2 + + i l h c i 1 + c - n o 8 8 2 n 9 9 6 ) 6 8 8 9 8 f 0 0 1 1 1 9 o 0 9 , , , 8 0 6 r r ) 0 s s 1 9 4 1 0 3 0 ) e e 1 e e 2 , 9 9 , 9 ) 4 5 9 6 ) g g r 1 c r h , 8 8 8 1 t 6 9 9 e 1 0 0 n n s 9 e ) n 6 9 t 1 1 t , 0 8 e e 1 g 1 , 9 0 1 g e 3 7 a r 3 8 r l l , , r r 9 1 , n 5 1 9 , n e r r 4 1 9 e 1 u u e l r r 7 e 6 1 , 9 8 , 1 e M l e e ) b 9 ) 9 1 g , l o o g r l e r u 1 0 , 5 r 1 , o g h t 0 o 1 4 1 9 , n u e 8 r n e u n t r c 9 B B e 9 9 e , 9 n 0 e P , 1 , 5 n e e o h 9 g R e c l o h n l l 1 g r i 1 l 1 e t 0 l 9 9 9 1 2 l g 8 s e c n l g l ü B e u s , n d , u B , n 9 O 1 1 1 i o n a e r 1 u r k u n ( o e g s o a a o o l ü s t 1 2 G s , , e e l , e c t t d e , P r o e r n P l r s u l u a r , 8 s B n e g B u g a a n G e o i l e e h r e u B u e t t i o e u n e 9 l t a n o i n x a e a u g H g t c c o e B n g h o a a 1 r e u e B n m c t u c n t n a B m n l l n n c k a i n ( s e m , t B o e B o u a i i a e h S f e i o u c u e u u ( t e i n e a ( i u l i l ( a i y i l c M n a s B t o s o p p c g n m d s a r u c u ( i a r n n n e y u i i i a n i s i M y n a o p t g t t r o o B B o d o o a a a o p s g n i l h i l l o y h s e b a n o l e c n d s e g g B e s o u n n o l p l u u B n l c B e m i i i u a G i t ( a ( i o n n c n a n a n a o m e p t a s a e e n i a i m m s h c i a t s i c i t a a a o r u d i t l g t n i r r a s s s s s s i a i t t e . . a e i r l t t s m c S e e r b p e n s f f l f g s i u u u u u u s g i d b p i e o s s i f f l l l l l l r i a e a o e r a v s e s t e n a s n a s s l k y i p p t a a u e e e e e e e r e o l e d s e s y u b s a a a s a u u r o u l h i b b b b b b p a d i t s r s s s s s s l u a f u a e h t r u e r r d m i u i i i i i i r a r t p e e t r m n e i g d r t t t t t t u g c a d h m m m m m m d r g g r t a e k i s l a i s h e i e a a i a e a o t i c n k n n n n n e e e e e e t n e d h e a a i t p t e t e c i c a t i t e l s n d c c c c c c n n l d o o o o o o o t a m m s i p y a b b o r l p i i p i a e p l o e o u s t a a a a a a i a a i d d d d d d o m y y h o t t i r p s d s s s o a t t t t t t a p n y r h p c t b r o r i g t s l o c o o o o o s a h h e e e e l s s s s s s p r t c r c i t t e m t t t t a o m e a u n a n o a n o n n n n n a m i a a a a a a a l l y l i l h a a u r c a r a a o a i a a a a a y y y y y y a P F P M A B A C A C R C L C B B D C T M M S M S S S S S L P L L L L L M M M M M M M Fishes from the Congolese coast of Lake Tanganyika 207 as until now C. sp. “bifrenatus” was only known from the scrutiny is needed to verify whether different populations central Tanzanian coast, and as Konings (1998) argues from the north-eastern shore are conspecific. that C. sp. “bifrenatus” could be a geographical morph of The species listed as T. moorii corresponds in coloura - C. popelini occurring on the opposite shoreline. As the tion to what Schupke (2003) calls the rainbow Tropheus . distribution of C. sp. “bifrenatus” seems much larger, and However, these specimens have five anal fin spines as we found no intermediate forms between the west coast instead of six in the “true” T. moorii from the type locali - C. sp. “bifrenatus” and C. popelini , this claim seems unlikely. ty in the extreme south of the lake. At the southernmost Given these new data and the disjointed and patchy distribu - locality of this survey, both T. moorii and a specimen with tions of Julidochromis spp., the relationships between all rep - six anal fin spines and with a clearly different colouration, resentatives of Chalinochromis and Julidochromis , and their corresponding to what Konings calls T. sp. “red”, were affiliation with Telmatochromis Boulenger, 1898, should be caught. Although the ranges of T. sp. “red” and T. moorii revised (Poll 1986, Konings 1998). Sturmbauer et al. (2010) are known to overlap near the village of Kiku (Fig. 1), this demonstrated the paraphyly of Julidochromis with respect to zone of sympatry is assumed to be just a few hundreds Chalinochromis , confirming the morphological tree of of meters long (Konings 1998). Yet as we found a speci - Takahashi (2003) in which representatives of Chalinochromis men, clearly belonging to T. sp. “red”, 60 km further and Julidochromis cluster together. It hence seems likely that north, this indicates that both species may occur in sym - the taxonomy of these genera should be revised. patry over a larger section of the coast. As members Cases of sympatry in Tropheus . Species of the genus of both species are known to mate assortatively (Egger et Tropheus are stenotypic rock-dwelling cichlids occurring on al. 2008), species boundaries are expected to be main - Lake Tanganyika’s rocky shores. Their limited dispersal tained in this contact zone. ability gave rise to over 100 mostly allopatric colour morphs. Species shifts along a north-south axis. A clear shift Although the genus’ taxonomy is incomplete and confusing, in species composition was observed along a large section most colour morphs can be classified in a few lineages of the western coast visited, especially along rocky (Schupke 2003) or presumed species (Konings 1998) mostly shores. For Tropheus, Simochromis, Eretmodus, supported by molecular studies (Sturmbauer et al. 2005 , Chalinochromis , and Julidochromis , this was discussed Egger et al. 2007). The distribution of these lineages reflects above, but the phenomenon was also noticed for some geological events in the lake’s history (Baric et al. 2003). other rock dwelling , with Neolamprologus In the present survey, representatives of at least six biolog - niger and N. toae being found in the northern half of the ical species were found along the western shore of Lake lake and Variabilichromis moorii in the southern basin. Tanganyika. As the number of anal spines is a taxonomi - The distributions found for the ectodine Ophthalmotilapia cally important character in Tropheus (see Brichard 1989, Pellegrin, 1904, with O. heterodonta found in the north - Snoeks et al. 1994), species delineation was based on anal ern and central part of the lake and O. ventralis found only spine counts as well as on colouration. As such, the same in the southern part correspond with Hanssens and Snoeks classification was obtained as in Konings (1998), whose (1999). These distributions reflect the historical split nomenclature is followed here. Both T. duboisi and of Lake Tanganyika into several subbasins (Snoeks 2000). T. annectens could easily be identified given their unique Yet, the many taxonomical problems that remain ( cfr. morphological features within the genus. The species list - supra ) indicate that the complexity of Lake Tanganyika ed here as T. aff. brichardi are characterized by their yel - cichlid systematics has been grossly underestimated low paired fins. They correspond to the nominal species (Snoeks 2000). Therefore, it is expected that such patterns described from the opposite shore, in having a modal anal will be observed once other taxa whose distribution spans spine count of six and in showing clear sexual dichroma - more than one subbasin, are revised, such as the rock- tism. In this species, females keep the juvenile pattern dwelling tropheine Petrochromis , with several unde - of vertical bands, whereas adult males obtain uniformly scribed species (Makasa and Snoeks 2003). dark green or brown flanks (Konings 1998). Where this Perspectives. As many localities visited during this species coexists with T. annectens , its territory is situated survey were sampled for the first time, many records are in the deeper part of the littoral zone, the upper part being new. For Simochromis pleurospilus, Altolamprologus occupied by its larger congener T. annectens . A similar sit - calvus , and Tropheus sp. “red”, the localities presented uation of T. annectens in sympatry with a genetically and here contain the northernmost localities where these phenotypically different Tropheus sp. was described in this species have hitherto been observed, while for region by Baric et al. (2003) and Sturmbauer et al. (2005). Simochromis marginatus, Julidochromis regani and The species listed here as T. sp. “black” is an undescribed Chalinochromis sp. “bifrenatus”, these results drastically species that probably deserves specific status as it is sym - enlarge their known distributions. For A. calvus , voucher patric with the species classified here as T. moorii as well specimens were until now only available from the south- as with T. annectens at the east coast of the lake, south western part of the lake, from Lukushia Bay to the of the Mahale Mountains (Konings 1998). As Schupke extreme South, whereas for S. pleurospilus , no collections (2003) splits this northernmost “species” in separate line - have been made outside of Zambia. Our records therefore ages, and as different mitochondrial lineages are known to show that these species occur more than 100 km further occur in this part of the lake (Egger et al. 2007), further north, which agrees with Büscher (1998) who found both 208 Van Steenberge et al. 3 8 + + + + + + + + + 2 e l b 7 + + + + + + + + + + a 2 T 6 + + 2 5 + + + 2 4 + 2 3 + + + + + + + + + 2 2 + + + + + + + + + + + 2 1 + + + + + + + + + + + + + + 2 0 + + + + + + + 2 9 + + + + + + + 1 ) 3 8 0 1 0 2 7 ( + 1 i h 6 s e t 1 a i h s a 5 g + k 1 n a i l T 4 p + + + 1 g m n a i 3 S + + + + + + + + 1 w o l 2 l + + + + + + + 1 o f 1 e + + + + + + 1 b i r t 0 + + + + + + + 1 y b 9 + + + + + + + d e i f 8 i + + + + + + s s a l 7 + + + + + + + + + + + c , a 6 + + + + + k i y 5 + + + + + + + + + n a g n 4 + a T 3 + + + + + + + + e k a 2 + + + + L n i 1 + , s e 7 h s 0 i 0 f 2 d , i i l r h o c i H 3 c t 0 e f 0 s o 2 l l ) , e i s ) 9 c ) B k 6 0 n ) 8 ) e , 0 9 ) i e 9 9 8 o 9 1 r 9 h 9 8 9 n r 1 , s ) 9 8 r 1 8 S ) , u a 9 ) 8 e r 1 , 1 c 6 t 8 h 4 r 8 1 e ) n , , c e ) 0 a 9 e r 9 9 r , g 8 h ) 8 9 k r 8 e ) s g 6 e 1 8 c n O 9 ) 9 a e 9 1 1 g 4 n n 0 g 9 , 1 a e 8 0 1 l l g T ) 8 , , e e n 9 9 n ) 8 , d l 9 1 r r l 0 9 s u r n , 4 e 1 e 8 1 9 9 n o 9 e , e l s u l 9 y e e 1 i o 7 , ) 4 r 1 , 4 ) 1 l l 8 g g P u r n o u 1 e r g , l 9 e 6 7 , ( ) 9 B 8 , t u r 1 e n n a o e o , r e n ( 1 g B ) 5 9 6 1 7 d r e e o e , g 6 S ( e g e a h B l r i l , H n r 9 B 9 1 ( e 5 l , 9 l g n 5 l ( ” h c n ( B l a e 2 e u s n u 4 1 , l g t 9 u 1 o s n e ( l s s l 9 e l S h s 4 g l o p s o o 9 , o l c n e l n 1 o , u o u u u 1 t l s c i l l u e 9 n u u t t 1 e s u l o P ü , B i h B l l o , P t g B l u d c u e l t a ( 1 o ( r a , l u o ( o ( i P l l o a a n l l s u h G n o B a n , i ) c u s i l i t B B P ( u o i h u ( l . B P s i g o o a u l e n s ( ( d n l n B o ( n l f i o d p b r P u n r L P a r e e s ( g e B i m i o t f n r e ( a o P o e n ( t a r s r i i o e ( a i B y a u i i i ( r t p c c a l u a P g c f r t d r l p u o r r c s i h i b h i i s m p e c a u a a h o i i i e e o e v n e n r e i p c c “ c c u t r e m l t c t i p r i n n s s s s s a r g s o p o o t l p c . i l a e i i e r o a m p i a a t u u u u u o r u e a e o r r e h r i p t a m c c e t t f m n t p b f l l l g g x g g g g g s p b a t d b s o l y s s s e m e s s s s s s s s s s l n o o o o o l e i s s s a s s s i r m l l l l l i e u u n u u u u u u u u u u i i i a i i i o i i c l i n t o o o o o c m g g s s g g g g g g g g g g p r r r r r n n m m m i i m m m e n s s n s s o o o i i o o o o o o o o o o i p p p p p i i o o o l l s o o o p l l l l l l l l l l o r u u m m g g i r r r r r r s o o o o o o o o o o o o h g g m m m m m m m m o o s o o h h h r r h h h r r r r r r r r r r c m l l r r o o i a a a a a d e o o o c i c c c c c p p l l p p p p p p p p p p l l l l l o o o r r l h h r n n u o o o o o o i r o o r o o o o o r i c c h h m m t t t m m m m m m m m m m a h n i i i i r r i i n n n b h c c i p o o p i i i a a a a a a a a a a a a a a a c d d d d p p d t p c e a l l l l l l l l l l l l l l l o o i i i i d d i i o i i n a o b m m m m a a a o m o o o o o o o o o o o l e e m m r p p p p p l l l l i l l l t t r r o a e e e e e e e e e e h h h a i l l y r e e e e a a e a y e e e u u C T V T L T T N N N N N N N N N N L L L T L J J L L L C O T T O C C A L A Fishes from the Congolese coast of Lake Tanganyika 209 3 8 + + + + 2 e l b 7 + + + + + + + + + a 2 T 6 + + + 2 5 + + + 2 4 + + 2 3 + + + + + + 2 2 + + + + + + + 2 1 + + + + + + + 2 0 + + + + + + + + + 2 9 + + + 1 8 + 1 7 + + 1 6 e + + + t 1 i s 5 g + + 1 n i l 4 p + + 1 m a 3 S + + + + + 1 2 + + + + + 1 1 + + + + + + + 1 0 + + + + 1 9 + + + + + 8 + + + + + + ) n o 7 + + + + + + + i t a u 6 + + + + + + + + n i t n 5 + + + + + + o c ( 4 3 + + + 2 + + + + + + + 1 ) 9 2 9 6 9 9 1 ) 1 ) , 9 , 2 a ) 9 s ) 6 y ) 6 e 8 e ) 6 9 a 8 0 h 1 g ) 6 ) t 0 1 k 9 9 , a t 8 e 0 a ) 9 , r ) 8 6 ) 1 a p ) 9 s g 9 ) e 4 1 8 1 0 9 N ) , 6 8 e a s 1 8 g 1 r , 9 M 9 , 9 9 6 t r 5 1 h p u r , 9 e n 8 9 8 9 t t 1 8 ) e r 0 e 9 , e o t e 8 g 9 1 s e 1 8 r e , i 1 1 l i 9 g 1 a g 8 1 n r 8 , u e , l 1 g v , 0 u h 1 n r , n e l e r 9 , r 1 o g s l e , n l 9 e o r , e 6 M i e e l o g 8 r e r l , r 9 a l n e 8 6 e u 1 g 0 t v r g e B l o n 1 g p e P u 9 e h u 9 5 g o , ( e e e n 9 ( l e n g u , n P g o r a 8 r o 8 l 9 n e 3 r g 1 l ( e u s B n e o e n l m k 1 l a p l e 1 i u B 1 l e ( 4 n , B e o t l e s 8 g a u o , o r B u l ( , p o , u g l e 9 s r r n ( u r u l 4 n a l o T e B i e o u P m f o n u t 1 l s e B ( o l l e e o ( 9 u g i s o o e s B l m o o , o B g l B d a o g l n r 1 s s o ( l n u B ( u ( u g a d r i r f B n l P a l o B n u ( t o y e , i e t i t i s B c o r r i i ( e l l t p r o i e o a n i u g t t t p l l a n n s y i l p s e e e i u i s l B P s e t a a e a l f e n r d s u o o p B i n i u o c m ( l o i i i n s e a g v e u p o n o o e s x o P p h t i i o n c i o a i s r o r v h n o i B r a n t e g a a a r r i p i t a B g i c s B l f r u c n l w m c m t x c o y u i u r i n a a a n s n i p e o i m u f e a r a e i i i a e o x e r n h n e o l t e o i l r o l t p m o t i l i r h d i p p p o l r n o m x m n m p m p e t c m s r n f e c a r i c i n a a a h i c a i a n n n s e r n s h i l l l s s s a i ( a e e n s o b l m c c i i i i i i y e i i i i a o o u f m o m c r r t t t p o i i m s r o t t p n p n d d n c a a a a a o s m s s m m m o o o s e p i r o s i i i i i n a ( o r s o a o o a t i e e o t o o o e a r u d t l a p p p p p h i i r t t e d e x m m m x s r r r r i l a e d a g l l l d g h a a a a a p c i t s a a g n u a n a n u h h h h l l l l l t d b o t i a a a t i c o n n o o a o i i i i i m u b b n n c d c c c i i s o s a y t t t t t i e o h h h o i d d h n y y e d r h s o t o o o l l t t t l s e r n m t r h l o o o o o h i l l l o o i t o h h o c p h h h p p p n b l n l l t u t a a t l p p t t u n n n n n r t r s i e a p p p a r y y o c a u a a y c y y a o e e e e e e c u l a a o e r e C T H X X X X E O X O O L H P G E C C C C C C E A C C C P B B B B B P 210 Van Steenberge et al. 3 8 + + + + + + + + + + + + 2 e l b 7 + + + + + + + + + + + a 2 T 6 + + + + + + + + + + 2 5 + + + + + + + + + + 2 4 + + + 2 3 + + + + + + + + + + + + + + + 2 2 + + + + + + + + + + + + + 2 1 + + + + + + + + + + + + + + + 2 0 + + + + + + + 2 9 + + + + + + + + + 1 8 1 7 + + + + + + + + 1 6 e + + + + + + + t 1 i s 5 g + + + + + + 1 n i l 4 p + + + + + + 1 m a 3 S + + + + + + + + + 1 2 + + + + + + + + + 1 1 + + + + + + 1 0 + + + + + + + + 1 9 + + + + + + + 8 + + + + + + + + + + + ) n o 7 + + + + + + + + + + i t a u 6 + + + + + + + + + + + + + n i t n 5 + + + + + + + + o c ( 4 + 3 + + + + + + + + + + 2 + + + + + + + + + 1 + + 5 7 9 1 , e d r e a n e d u A n 0 e ) 6 d e 9 g 1 n 3 a ) a , 8 p 2 s ) ) v 9 ) 4 9 a 4 8 8 9 s 5 4 8 ) 1 s 1 8 v 9 7 9 ) 9 u 9 9 9 8 9 y , 9 ) 0 a 1 9 8 4 8 o 1 8 8 9 a 1 h 8 4 0 i 7 , 1 1 4 1 , w 1 1 8 0 k , l ) 1 T 9 9 v 2 , s r , 8 e 9 l , , , 1 0 o 6 r e 4 , 8 1 t r r 9 e e r r n 4 1 o r a r , 9 5 e e e 9 1 , 1 e e e h g T r 8 9 , e P r p 1 9 t g g 8 s l , 0 , h g m e n ( t t 9 n 1 g e l r t , 1 s n ” n 1 a e 5 n n 9 e g a e r i 8 , n e g s o l m i n e , e h l l , e s 9 5 n e l Y l e l r t 1 s h n n l r l u o P ü e s l l M t r e 1 9 g u r e ( u g e , i e o o s u l o o u l r l f n o , G 1 n r N e o s h o i h u o u P l f l ( e e t o u P B t ü n e , l u i l n t s n h B o l B r g o B “ n e t v B o e ( s e h s N a a ( G u o e u l t n B r ü a l i a ( i s s B u u i i P P i i t ( o e n e l m s t a t i u r M n i u u l d r i G a p a o e n t t a a n a c s B n t e r ( g n m u a s l ” u r l f a c c v d a o n g u n a t f o s i l o a i i s o i o o t k n u c a i o t d t t s o r e i r e h i e M r u ” n c g a r a f s s t y B a c r o h c d c g i w r u s i e m l a d s t s r i u b p o o r n m s i i t a o l s e a e i b a e e o r o r i r i h b i l a n n a c o r a a i i h r b s r r a t n m t o p m f f b i o e c m d p b a a d l c i “ “ s h y o s s o ( . i y y s s s s s s b n e i . . r c s s s s r m s f i i i i i i o u a i c c m i i i i u n p f e i p p e o m h o d m s t o n m m m m m m s m s d a a r i s s p c o m m m m s i o o r m o o o o o o o r h u u a n o s s s s s s i r d o o o o l u c d r r r r r r l i h c i i h r r r r s e d d u u u u u u i h n d o e h h h h h h c r t c o h n h h h h o o o e e e e e e c a o c c c c c c o h o o c o c c c c h a d h h h h h h o m l t h l t e o o o o m o o m n o t p o o o o t g u t t p p p p p p n r r r r r r p a p a b e b t t t t t t o t e a n e e e i o o o o o o m a m m m m a n r t s s e e e e e e r r r r i o r r r r r r i i i i p a T H A H T T T T T T S S S E E S P P P P P P P E G L L C T S T Fishes from the Congolese coast of Lake Tanganyika 211 species at the village of Tembwe (which is not to be con - (Perciformes: Cichlidae) in the Lake Malawi basin. fused with Cape Tembwe, a locality further north that was International Journal of Evolutionary Biology 2011 sampled in this study), just 10 km further south of our (835946): 1–11. DOI: 10.4061/2011/835946 southernmost sampling site (Fig. 1). Although littoral Baric S., Salzburger W., Sturmbauer C. 2003. cichlid communities are much better known than those Phylogeography and evolution of the Tanganyikan cichlid that occur in deeper water or further from shore, our find - genus Tropheus based upon mitochondrial DNA sequences. ings illustrate that our knowledge of their specific compo - Journal of Molecular Evolution 56 (1): 54–68 . DOI: sition remains fragmentary. Therefore, it can be expected 10.1007/s00239-002-2380-7 that sampling in less accessible habitats will lead to even Bell-Cross G., Bell-Cross B. 1971. Introduction of Limnotrissa more discoveries. Between 1989 and 1997, 12 new cich - miodon and Limnocaridina tanganicae from Lake lids were described by Büscher (e.g. , 1997), who collect - Tanganyika into Lake Kariba. Fisheries Research Bulletin ed all of them in the south-western Congolese part of the Zambia 5: 207–214. lake, mostly in habitats not targeted during this survey. Boulenger G.A. 1915. Catalogue of the fresh-water fishes This illustrates the need for more field surveys, especial - of Africa in the British Museum (Natural History). Volume ly along less explored coast and further from shore, as the III. Trustees of the BMNH, London. cichlid community at greater depths is completely differ - Brichard P. 1978. Fishes of Lake Tanganyika. T.F.H. ent from the one found at rocky shores. Publications, Neptune City. New insights in the distribution of many species, and Brichard P. 1989. Pierre Brichard’s book of cichlids and all the other the near-unavailability of such data in the published liter - fishes of Lake Tanganyika. T.F.H. Publications, Neptune City. ature, underpin the fact that, despite the large number of Brown K.J., Rüber L., Bills R., Day J.J. 2010. 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