Abstract
Five new rheophilic haplochromine cichlid species are described from the Upper Congo drainage of Zambia and the Democratic Republic of the Congo: Orthochromis mporokoso sp. nov. and O. katumbii sp. nov. from the Bangwelu-Mweru ecoregion, O. kimpala sp. nov. and O. gecki sp. nov. from the Upper Lualaba ecoregion, and O. indermauri sp. nov. from the Lufubu River of the Lake Tanganyika ecoregion. Orthochromis kimpala sp. nov, O. gecki sp. nov., and O. indermauri sp. nov. are distinguished from all currently valid species of the genus Orthochromis Greenwood 1954, except for O. torrenticola (Thys van den Audenaerde 1963), by the presence of eggspots or eggspot-like maculae on the anal fin (vs. no eggspots). The three species can be easily distinguished from O. torrenticola by having three anal spines (vs. four anal spines). Moreover, all five new species can be individually distinguished from all currently known rheophilic taxa placed in the genera Orthochromis, Schwetzochromis Poll 1948 and the rheophilic species of the genus Haplochromis Hilgendorf 1888 (e.g. H. bakongo Thys van den Audenaerde 1964, H. snoeksi Wamuini Lunkayilako & Vreven 2010, H. vanheusdeni Schedel et al. 2014) either based on meristic values, morphometric distances and colouration patterns, or on a combination of them.
References
Barel, C.D.N., van Oijen, M.J.P., Witte, F. & Witte-Maas, E.L.M. (1977) An introduction to the taxonomy and morphology of the haplochromine Cichlidae from Lake Victoria. Netherlands Journal of Zoology, 27 (4), 333–389.
https://doi.org/10.1163/002829677X00199Balon, E.K. & Stewart, D.J. (1983) Fish assemblage in a river with unusual gradient (Luongo, Africa Zaire system), reflections on river zonation, and description of another new species. Environmental Biology of Fishes, 9, 225–252.
https://doi.org/10.1007/BF00692373Boulenger, G.A. (1899) Matériaux pour la faune du Congo. Poissons nouveaux du Congo. Cinquième Partie. Cyprins, Silures, Cyprinodontes, Acanthoptérygiens. Annales du Musée du Congo, Series Zoology, 1 (5), 97–128.
Boulenger, G.A. (1902) Contributions to the ichthyology of the Congo.—II. On a collection of fishes from the Lindi River. Proceedings of the Zoological Society of London, 1 (4), 265–271.
Boulenger, G.A. (1905) On a collection of fishes from Lake Bangwelo. Annals and Magazine of Natural History, Series 7, 16 (96 Art. 72), 642–647.
https://doi.org/10.1080/03745480509443309Broadley, D.G. & Cotterill, F.P.D. (2004) The reptiles of south-east Katanga, an overlooked ‘hot spot’. African Journal of Herpetology, 53, 35–61.
https://doi.org/10.1080/21564574.2004.9635497Clabaut, C., Salzburger, W. & Meyer, A. (2005) Comparative phylogenetic analyses of the adaptive radiation of Lake Tanganyika cichlid fish: Nuclear sequences are less homoplasious but also less informative than mitochondrial DNA. Journal of Molecular Evolution, 61, 666–681.
https://doi.org/10.1007/s00239-004-0217-2Cotterill, F.P.D. (2005) The Upemba lechwe, Kobus anselli: an antelope new to science emphasizes the conservation importance of Katanga, Democratic Republic of Congo. Journal of Zoology, 265, 113–132.
https://doi.org/10.1017/S0952836904006193
David, L. (1937) Poissons de l'Urundi. Revue de Zoologie et de Botanique Africaines, 29 (4), 413–420.
De Vos, L. & Seegers, L. (1998) Seven new Orthochromis species (Teleostei: Cichlidae) from the Malagarasi, Luiche and Rugufu basins (Lake Tanganyika drainage), with notes on their reproductive biology. Ichthyological Exploration of Freshwaters, 9 (4), 371–420.
De Zeeuw, M.P., Westbroek, I., van Oijen, M.J.P. & Witte, F. (2013) Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania. ZooKeys, 256, 1–34.
https://doi.org/10.3897/zookeys.256.3871Dunz, A. & Schliewen, U.K. (2010) Description of a new species of “Tilapia” Smith, 1840 (Teleostei: Cichlidae) from Ghana. Zootaxa, 2548, 1–21.
Dunz, A.R. & Schliewen, U.K. (2013) Molecular phylogeny and revised classification of the haplotilapiine cichlid fishes formerly referred to as "Tilapia". Molecular Phylogenetics and Evolution, 68, 64–80.
https://doi.org/10.1016/j.ympev.2013.03.015Free Spatial Data DIVA-GIS (2018) Available from: http://www.diva-gis.org/Data (accessed 29 January 2018)
Greenwood, H.P. & Kullander, S.O. (1994) A taxonomic review and redescription of Tilapia polyacanthus and T. stormsi (Teleostei: Cichlidae), with descriptions of two new Schwetzochromis species from the Upper Zaire River drainage. Ichthyological Exploration of Freshwaters, 5 (1), 161–180.
Greenwood, P.H. (1954) On two species of cichlid fishes from the Malagarazi River (Tanganyika), with notes on the pharyngeal apophysis in species of the Haplochromis group. Annals and Magazine of Natural History, Series 12, 7 (78), 401–414.
https://doi.org/10.1080/00222935408656059Greenwood, P.H. (1979) Towards a phyletic classification of the ‘genus’ Haplochromis (Pisces, Cichlidae) and related taxa, part I. Bulletin of the British Museum (Natural History) Zoology, 35, 265–322.
https://doi.org/10.5962/bhl.part.20455Hammer, Ø., Harper, D.A.T. & Ryan, P.D. (2001) PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4 (1), 9.
Hilgendorf, F.M. (1888) Fische aus dem Victoria-Nyanza (Ukerewe-See), gesammelt von dem verstorbenen Dr. G. A. Fischer. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1888, 75–79.
Huysseune, A. Rüber, L. & Verheyen, E. (1999) A single tooth replacement pattern generates diversity in the dentition in cichlids of the tribe Eretmodini, endemic to Lake Tanganyika (Teleostei: Cichlidae). Belgian Journal of Zoology, 129, 157–174.
Indermaur, A. (2014) Über den Tellerrand geschaut - Buntbarsche aus dem Lufubu. Die Aquarien- und Terrarien-Zeitschrift, 10, 16–23.
Jorissen, M.W.P, Pariselle, A., Huyse, T., Vreven, E.J., Snoeks, J., Volckaert, F.A.M., Chocha Manda, A., Kapepula Kasembele, G., Artois, T. & Vanhove, M.P.M. (2017) Diversity and host specificity of monogenean gill parasites (Platyhelminthes) of cichlid fishes in the Bangweulu-Mweru ecoregion. Journal of Helminthology, 92 (4), 417–437.
https://doi.org/10.1017/S0022149X17000712Katemo Manda, B., Colinet, G., Manda, A.C., Marquet, J.P. & Micha, J.C. (2010) Evaluation de la contamination de la chaîne trophique par les éléments traces (Cu, Co, Zn, Pb, Cd, U, V et As) dans le bassin de la Lufira supérieure (Katanga/RD Congo). Tropicultura, 28 (4), 246–252.
Koblmüller, S., Schliewen, U.K., Duftner, N., Sefc, K.M., Katongo, C. & Sturmbauer, C. (2008) Age and spread of the haplochromine cichlid fishes in Africa. Molecular Phylogenetics and Evolution, 49, 153–169.
https://doi.org/10.1016/j.ympev.2008.05.045Koblmüller, S., Katongo, C., Phiri, H. & Sturmbauer, C. (2012) Past Connection of the Upper Reaches of a Lake Tanganyika Tributary with the Upper Congo Drainage Suggested by Genetic Data of Riverine Cichlid Fishes. African Zoology, 47 (1), 182–186.
https://doi.org/10.3377/004.047.0115Kullander, S.O. & Roberts, T.R. (2011) Out of Lake Tanganyika: endemic lake fishes inhabit rapids of the Lukuga River. Ichthyological Exploration of Freshwaters, 22, 355–376.
Matschiner, M., Musilová, Z., Barth, J.M.I., Starostova, Z., Salzburger, W., Steel, M. & Bouckaert, R. (2016) Bayesian Phylogenetic Estimation of Clade Ages Supports Trans-Atlantic Dispersal of Cichlid Fishes. Systematic Biology, 66, 3–22.
https://doi.org/10.1093/sysbio/syw076Meyer, B.S., Matschiner, M. & Salzburger, W.A. (2015) A tribal level phylogeny of Lake Tanganyika cichlid fishes based on a genomic multi-marker approach. Molecular Phylogenetics Evolution, 83, 56–71.
https://doi.org/10.1016/j.ympev.2014.10.009
Poll, M. (1948) Descriptions de Cichlidae nouveaux recueillis par le Dr. J. Schwetz dans la rivière Fwa (Congo belge). Revue de Zoologie et de Botanique Africaines, 41 (1), 91–104.
Poll, M. (1967) Contribution à la faune ichthyologique de l'Angola. Publicações Culturais, Companhia de Diamantes de Angola (DIAMANG), Lisboa, 75, 1–381, Pls. 1–20.
Poll, M. (1976) Poissons. Exploration du Parc National de l'Upemba, Mission G. F. de Witte, Bruxelles. Reports of the Swedish Deep-Sea Expedition 1947-1948, 73, 1–127.
Poll, M. (1986) Classification des Cichlidae du lac Tanganika. Tribus, genres et espèces., Collection in 8 - 2ème série; Bruxelles, 45 (2), 1–163.
Roberts & Stewart, D.J. (1976) An ecological and systematic survey of fishes in the rapids of the lower Zaire or Congo River. Bulletin of the Museum of Comparative Zoology, 147 (6), 239–317.
Salzburger, W., Meyer, A., Baric, S., Verheyen, E. & Sturmbauer, C. (2002) Phylogeny of the Lake Tanganyika cichlid species flock and its relationship to the Central and East African haplochromine cichlid fish faunas. Systematic Biology, 51 (1), 113–135.
https://doi.org/10.1080/106351502753475907Schedel, F.D.B., Friel, J.P. & Schliewen, U.K. (2014) Haplochromis vanheusdeni, a new haplochromine cichlid species from the Great Ruaha River drainage, Rufiji basin, Tanzania (Teleostei, Perciformes, Cichlidae). Spixiana, 37 (1), 135–149.
Schedel, F.D.B. & Schliewen, U.K. (2017) Hemibates koningsi spec. nov: a new deep-water cichlid (Teleostei: Cichlidae) from Lake Tanganyika. Zootaxa, 4321 (1), 92–112.
https://doi.org/10.11646/zootaxa.4312.1.4Schwarzer, J., Swartz, E.R., Vreven, E., Snoeks, J., Cotterill, F.P.D., Misof, B. & Schliewen, U.K. (2012) Repeated trans-watershed hybridization among haplochromine cichlids (Cichlidae) was triggered by Neogene landscape evolution. Proceedings of the Royal Society B, 279 (1746), 4389–4398.
https://doi.org/10.1098/rspb.2012.1667Thieme, M.L., Abell, R., Stiassny, M.L.J., Skelton, P., Lehner, B., Teugels, G.G., Dinerstein, E., Kamdem-Toham, A., Burgess, N. & Olson, D. (2005) Freshwater Ecoregions of Africa and Madagascar: A Conservation Assessment. Island Press, Washington, D.C., 483 pp.
Thys van den Audenaerde, D.F.E. (1963) Description d'une espèce nouvelle d'Haplochromis (Pisces, Cichlidae) avec observations sur les Haplochromis rhéophiles du Congo oriental. Revue de Zoologie et de Botanique Africaines, 68, 140–152.
Thys van den Audenaerde, D.F.E. (1964) Les Haplochromis du Bas-Congo. Revue de Zoologie et de Botanique Africaines, 70, 154–173.
Van Oijen, M.J.P., Snoeks, J., Skelton, P.H., Maréchal, C. & Teugels, G.G. (1991) Haplochromis. In: Daget, J., Gosse, J.P., Teugels, G.G. & Thys van den Audenaerde, D.F.E. (Eds.), Check-list of the freshwater fishes of Africa (CLOFFA). Vol. 4. ISNB, Bruxelles, MRAC, Tervuren and ORSTOM, Paris, pp. 100–184.
Van Oijen, M.J.P. (1996) The generic classification of the haplochromine cichlids of Lake Victoria, East Africa. Zoologische Verhandelingen, Leiden, 302, 57–110.
Vandervennet, E., Wautier, K., Verheyen, E. & Huysseune, A. (2006) From conical to spatulate: Intra- and interspecific changes in tooth shape in closely related cichlids (Teleostei; Cichlidae: Eretmodini). Journal of Morphology, 267, 516–525.
https://doi.org/10.1002/jmor.10418Wamuini Lunkayilakio, S. & Vreven, E.J. (2010) 'Haplochromis' snoeksi, a new species from the Inkisi River basin, Lower Congo (Perciformes: Cichlidae). Ichthyological Exploration of Freshwaters, 21 (3), 279–3287.
Weiss, J.D., Cotterill, F.PD. & Schliewen, U.K. (2015) Lake Tanganyika—A ’Melting Pot’ of ancient and young cichlid lineages (Teleostei: Cichlidae)? PLOS ONE, 10, e0125043.
https://doi.org/10.1371/journal.pone.0125043White, F. (1983) The vegetation of Africa. A descriptive memoir to accompany the UNESCO/AETFAT/UNSO vegetation map of Africa. Natural Resources Research No. 20. UNESCO, Paris, 352 pp.