There is a great variability and diversity of social and breeding behaviours in cichlids, and this may contribute to rapid speciation. An important aspect of understanding adaptive radiations is concerned with the mode of speciation which led to their diversification. Second, most genes we identify as being under positive selection have accumulated relatively few single nucleotide polymorphisms (SNPs) owing to the short divergence time between species [15]. When Nile tilapia, infected with P. anguilliseptica, were exposed to a bath of the C. alata at 100 ppm, fish mortality was eliminated completely. Nine of these EST pairs showed a strong signal of positive selection (Ka/Ks > 1; 75–100% of the full-length gene; figure 3). The most prevalent tooth type is curved, with a sharp point, and may be uni- or bicuspid. Nucleotide variation and substitution patterns were examined using the software package MEGA (Kumar et al., 1993; version 1.01). Mechanisms of species divergence through visual adaptation and sexual selection: Perspectives from a cichlid model system. This book covers the history of these fishes over the last 250 million years by exploring the links between their biological evolution and the paleogeographic and environmental transformations of our planet, whether these be gradual or ... Unlike complex traits such as body shape, across many vertebrate taxa coloration tends to be of a simple genetic basis and therefore a more tractable target for comparative genomics [35,36]. Exploring the Expression of Cardiac Regulators in a Vertebrate Extremophile: The Cichlid Fish. Tilapia hybrids (particularly Blue–Nile hybrids) have played a significant role in global tilapia production since mid-2000s, contributing 6.7% in 2017. CiteSeerX - Document Details (Isaac Councill, Lee Giles, Pradeep Teregowda): Abstract: An Agent-Based model of speciation in cichlid fish has been implemented. The models are based on a combination of laboratory experiments and field observations on the ecology, male and female mating behavior, and inheritance of sex-determination and color polymorphisms. The behavioral and physical changes resulting from intense speciation in . In the polygynous mating system of A. burtoni, social dominance and therefore reproductive potential is associated with differences in gonad size, growth, hormone levels and coloration. In the upper pharyngeal jaws, the three dentigerous plates may carry different types of tooth. In 2011, Lake Malawi-type cichlids were discovered in a series of volcanic crater lakes in southern Tanzania, so this process might be happening in several places at once. Mol Ecol. For this species, Nagl et al. 435 - 443 View Record in Scopus Google Scholar Photo credit: P. Eriksson. This volume explores the phylogeny of fishes at multiple taxonomic levels, uses methods of analysis of molecular data that apply both within and between fish populations, and employs molecule-based phylogenies to address broader questions ... (three-striped damselfish), TABLE III. ichlidae), Oxidative phosphorylation gene transcription in whitefish species pairs reveals patterns of parallel and nonparallel physiological divergence, Evolution of genomic structural variation and genomic architecture in the adaptive radiations of African cichlid fishes, sixteen lacustrine and riverine habitat cichlids. Epigenetic variation can alter transcription and promote phenotypic divergence between populations facing different environmental challenges. To the best of the author's knowledge, the data set includes all available ray-finned fish taxa or their representatives from well-corroborated clades (e.g., Cypriniformes, Cichlidae) that have complete cytochrome b gene sequence data. Extensive MHC variability in cichlid fishes of Lake Malawi has been reported (Klein et al., 1993). Can male-male competition stabilize speciation? The low number of genes under positive selection between A. amarillo and A. sagittae (nine out of 1612 ESTs, or 0.6%) agrees with the previous findings from Lake Apoyo Midas cichlids, wherein 0.8 per cent (14 of 1721) of shared ESTs were found to be under selection [75]. Proceedings Biological sciences The Royal Society. Of 11 candidate genes for ecomorphological differentiation in African cichlids, none showed signs of selection between A. amarillo and A. sagittae. 2014). [85]). Nature 455:620-626. light spectrum photo by Dan Young and used under a Creative Commons license ; cichlid photo by Erica Marshall of muddyboots.org and used under a Creative Commons license for the basal haplochromine Astatotilapia burtoni [8–11]). Some studies on this direction are in progress in salmonid species (K. M. Miller, pers. Cichlid fishes are spectacularly species-rich and phenotypically diverse in morphology, behaviour and coloration, and therefore have become a ‘non-model’ model system for studying genomic diversification by natural and sexual selection [1–3]. Very rapid speciation occurred when cichlid fishes were isolated in Lake Victoria. Please enable it to take advantage of the complete set of features! Figure 2. Cichlid fishes are spectacularly species-rich and phenotypically diverse in morphology, behaviour and coloration, and therefore have become a 'non-model' model system for studying genomic diversification by natural and sexual selection [1-3].Many of the more than 2000 species have diversified based on ecological niche within lakes . The levels of these parameters decreased significantly in the blood of fish treated with pathogenic bacteria. -, J Mol Evol. Prevention and treatment information (HHS). In the rivers: Multiple adaptive radiations of cyprinid fishes (Labeobarbus) in Ethiopian Highlands. Figure 1.9. A single gene was found to be associated with the OB phenotype: Pax7 expression is increased in tailfins of OB individuals in all three Lake Malawi species examined though no sequence differences were found in the Pax7 coding regions [56]. The Cichlid Speciation Project (CSP) is an ALife simulation system for investigating open problems in the speciation of African cichlid fish. 15, and Unh011, Unh012 and Unh013; ref. As result, the search for alternative, unconventional and more environmentally friendly disease treatments and control strategies is increasing. Speciation through sensory drive in cichlid fish. Theme Issue 'Patterns and processes of genomic divergence during speciation' compiled and edited by Patrik Nosil and Jeffrey L. Feder, Genomics of adaptation and speciation in cichlid fishes: recent advances and analyses in African and Neotropical lineages, Candidate genes of African cichlid species related to vision and light perception, bone and tooth development, and morphology. TEC counts and lymphocyte counts of infected fishes previously fed with control diet decreased significantly (43%, 36%), whereas it was minimal in fishes (21%, 8%) fed a vitamin C–supplemented diet. What evidence suggests that the Central European blackcap lineage is beginning to split? And the object lessons are—you guessed it—cichlid fish. Accelerated evolution and a cichlid-specific isoform of the pigmentation candidate gene mitf were also suggested as relevant for the rapid evolution of different colorations [50]. The second edition of Bones and Cartilage includes the most recent knowledge of molecular, cellular, developmental and evolutionary processes, which are integrated to outline a unified discipline of developmental and evolutionary skeletal ... Only further genomic and transcriptomic comparisons between Neotropical and African cichlid adaptive radiations, ideally aided by mapping and whole genome comparisons, can discern these differences. Predatory fish were added. However, commercial tilapia culture is currently restricted to about 10 species (Table 1.1). To address such questions of intra- and inter-adaptive radiation genome evolution, we analysed newly generated ESTs for two sympatric, endemic, ecologically divergent, very young species of Midas cichlid from the Nicaraguan crater lake Xiloá (ca 6 kyr old; figure 2). The oral dentition of Neotropical cichlids shows much less variation than in their African counterparts (Casciotta and Arratia, 1993). Copeia 2014:556-560. pdf. These findings are similar to those found in functional mammalian class II genes, thereby implying that the function of these genes is probably the same as those of the mammalian MHC. African cichlid fish: a model system in adaptive radiation research. Across African cichlid radiations, sexual selection on colour pattern is one of the most important forces for speciation [42], suggesting that positive selection may be acting on the gene(s) responsible for coloration. Pseudomonas fluorescens were found to cause chronic mortality in farmed Nile tilapia in Japan (Miyashita, 1984). To analyse genomic diversity and selection in a Neotropical radiation, we conducted a comparative transcriptomic analysis between sympatric, ecologically divergent crater-lake Midas cichlids (Lake Xiloá Amphilophus amarillo and Amphilophus sagittae). Discussion and extension questions. We review here recent molecular data on population differentiation and phylogenetics, which have helped to unravel, to some extent, the patterns and processes that led to the formation and ecological maintenance of cichlid species flocks. This lake harbours about 250 species of cichlid fish, which are highly diverse in terms of morphology, behaviour, and ecology. A news story on speciation in cichlid fish. Blotched phenotype in general is correlated with increased aggressive behaviour [52] and associated with sexual selection by male preference in Lake Victoria OB and WB species [43] and Lake Malawi OB species [49], such that only in polymorphic populations do males show a preference for blotched females. African cichlid fish: a model system in adaptive radiation research. Epub 2008 Dec 16. 2003 Sep;12(9):2497-504 Population divergence along the benthic-limnetic axis is common in freshwater fish (Robinson and Wilson 1994) and is the basis of sympatric speciation in at least two radiations of Crater Lake Midas cichlids (Barluenga et al. Infected fish suffered from mass mortality (up to 100%), erratic swimming and skin darkening. Therefore, this colour pattern may be a mechanism of rapid sympatric speciation by sexual selection [53–55] and represents a simple genetic basis of behavioural phenotypes. From: The Egyptian Journal of Aquatic Research, 2021, Barry Berkovitz, Peter Shellis, in The Teeth of Non-Mammalian Vertebrates, 2017. This species pair, Amphilophus amarillo and Amphilophus sagittae, diverged along a benthic–limnetic phenotypic axis [15,76], similar to the Midas species pair from the older crater lake Apoyo [4,75]. erichthys bartoni While this type of speciation may not be as common as the other types, it does occur, as evidenced by the sympatric speciation of cichlid fish in Africa and Nicaragua. As part of the 2017-2018 Fellows' Presentation Series at the Radcliffe Institute for Advanced Study, Axel Meyer RI '18 shows us why fish are much more intere. V. 273 (1597); pp. Specifically, the relative importance of intrinsic biological characteristics such as ecology, anatomy, and behavior versus abiotic factors such as geological history, geographic structuring of the lake basin, barriers to gene flow, and fluctuations of the lake level is controversial. They support more diversified ichthyofauna than the previously described lakes. OB and WB patterning (figure 1b), while melanin-disrupting and female-linked in all species tested for the radiations in lakes Malawi and Victoria, are determined by different genes [51]. Other transcriptomic approaches that can infer abundance (e.g. Both of these species are unique to their respective lakes and are stream spawners, migrating into FW rivers for breeding and returning to the lake for feeding and development. Research on speciation in cichlids has to date primarily focused on the species-rich flocks of the African Rift lakes. Global production of major farmed fish species (t, %) in 2017. Download. The genetic basis of this behavioural variability and plasticity has recently been a focus of research using genomic and transcriptomic methods. We suggest four possible reasons for this finding. Photo credit: J. Desjardins. (a) A breeding pair of Midas cichlids (Amphilophus xiloaensis) from crater Lake Xiloá, Nicaragua. Photo credit: P. Eriksson. Here we use African cichlid fish to compare the inviability rates of F2 hybrids with rates found in F1 crosses and the corresponding grandparental-type crosses. Moreover, these two sex chromosomes are not overlapping with the sex chromosomes of the Nile tilapia (O. niloticus), which are located at different chromosomal regions [59]. Mechanisms of rapid sympatric speciation by sex reversal and sexual selection in cichlid fish Genetica , 112-113 ( 2001 ) , pp. There are two very distinct ectomorphs or forms in the population: a yellow-green one that lives along the shore, and a blue-black one that lives at the bottom of the lake. Libraries were normalized in order to maximize the total length and number of ESTs sequenced, though this comes at the cost of gene-expression inference. P First book to outline the fundamental principles of social evolution underlying the stunning diversity of social systems and behaviours. understand Cichlid fish speciation, rather than the ecology of the lakes. Found inside – Page 206It is likely that the combination between intersexual and intrasexual selection for male nuptial coloration further accelerates reproductive isolation and rapid speciation of cichlids in Lake Victoria. Cichlid fishes in Lake Malawi are ... Table 1.Candidate genes of African cichlid species related to vision and light perception, bone and tooth development, and morphology. Darwin's Dreampond blends a discussion of the principles of neo-Darwinian evolution and speciation with a history of Lake Victoria's ecosystem. The book was shortlisted for the BP Natural World Book Prize. Cichlid fish from Lake Apoyeque, Nicaragua, show evidence of sympatric speciation. 2002 Jun;19(6):865-83. doi: 10.1093/oxfordjournals.molbev.a004144. The ratio of non-synonymous to synonymous substitutions (Ka/Ks) was estimated with maximum likelihood in PAML v. 4 [79]; pairs with Ka/Ks > 1 were confirmed by outgroup comparison with African cichlids. There may be considerable sex-specific and species-specific gene regulation associated with breeding systems such as monogamy or polygyny [63]. The effect of probiotics on four hematological parameters, viz. In addition to the actinopterygians, DNA sequences for cytochrome b were obtained from GenBank for the sharks Carcharhinus plumbeus (Martin et al., 1992; GenBank L08032) and Galeocerdo cuvier (Martin et al., 1992; gb L08034) and for the following tetrapods: African clawed frog, Xenopus laevis (Roe et al., 1985; gb M10217), opossum (Monodelphis domestica, Ma et al., 1993; gb X70674), human (Anderson et al., 1981), and pygmy right whale (Caperea marginata, Arnason and Gullberg, 1994; embl X75586) to serve as outgroup taxa. Therefore, research efforts have been concentrated on optimizing production with ecofriendly alternatives to the therapeutic use of antimicrobials. Cichlid fish in Lake Victoria may serve as yet an example of sympatric speciation. Coding regions of the putative orthologous genes were annotated by comparison with currently available vertebrate proteins. Prevention of the disease with the help of chemical compounds may prove harmful for fish and the end consumer. In this case, even comparing parallel adaptive phenotypes across lineages would still identify non-overlap of genes under selection because the underlying genetic processes and architecture of parallel phenotypes might be entirely different. As of early 2011, the tilapia genome (Oreochromis niloticus) was completed and assembled, and whole genome sequencing of four other African Rift Lake species is near completion (F. Di Palma 2011, personal communication). Moreover, some genomic regions contain multiple quantitative trait loci (QTLs). These tiny lakes may hold the key to understanding cichlid speciation, but they are extremely fragile habitats. 2006; Elmer et al . See §1b,c for more details. Kullander recognises the following Neotropical subfamilies: Astronotinae (Astronotus, Chaetobranchus), Geophaginae (for In the 1950s, an irrigation dam was built on the stream . This site needs JavaScript to work properly. Lake Tanganyika is the oldest of the Great Ancient Lakes in the East Africa. Food fisheries have existed for both species although both are threatened by population declines due to overfishing. Although this form of speciation may be observed in a variety of species, including the apple maggot fly, bacteria, and cichlid fish, it is difficult to determine when it happens spontaneously. This image shows two helpers-at-the-nest (left) and a breeder female (right). Frequently, many closely related species coexist that have diverged without geographical isolation and therefore presumably, or demonstrably, under divergent selection and through ecological speciation [4,5]. In the 1950s, an irrigation dam was built on the stream . In a series of twenty chapters, Ernst Mayr presents a consecutive story, beginning with a description of evolutionary biology and ending with a discussion of man as a biological species. Furthermore, the number of species in cichlid radiations increases with lake size, supporting the prediction that species diversity increases with habitat heterogeneity, but also with opportunity for isolation by distance. Saccharomyces cerevisiae, Transcriptome sequencing of three Ranunculus species (Ranunculaceae) reveals candidate genes in adaptation from terrestrial to aquatic habitats, Preface: Speciation research in ancient lakes – classic concepts and new approaches, The Evolutionary Genomics of Cichlid Fishes: Explosive Speciation and Adaptation in the Postgenomic Era, Genomic architecture of ecologically divergent body shape in a pair of sympatric crater lake cichlid fishes, Sexual selection and genetic colour polymorphisms in animals, A Hybrid Genetic Linkage Map of Two Ecologically and Morphologically Divergent Midas Cichlid Fishes ( What, if anything, is a Tilapia?-mitochondrial ND2 phylogeny of tilapiines and the evolution of parental care systems in the African cichlid fishes. Provides descriptions of the methods and tools used in molecular population genetics, which has combined advances in molecular biology and genomics with mathematical and empirical findings to uncover the history of natural selection and ... By continuing you agree to the use of cookies. Table II lists the 31 actinopterygian species examined in this study and their current classification (Nelson, 1994). SWS1 harbours the greatest diversity of functionally critical sites compared to other opsin genes in Malawi cichlids [. Discrepancies between the restriction fragment length polymorphism (RFLP) pattern of mtDNA and the species tree based on morphological characters, however, led Moran and Kornfield (1993) to suggest an ancestral polymorphism in the founding populations of the Lake Malawi flocks, which hinders an accurate determination of their phylogenetic relationships. Mechanisms of speciation in cichlid fish were investigated by analyzing population genetic models of sexual selection on sex-determining genes associated with color polymorphisms. In this study, three closely related species of the Pseudotropheus (now Maylandia) zebra complex were studied with regard to their courtship sounds -, Proc Natl Acad Sci U S A. In a second (unpublished) approach, locus-specific PCR primers were used to amplify microsatellite repetitive elements to determine allele size frequencies among cichlid species from Lake Victoria. Lake Tanganyika's cichlid diversity has evolved through explosive speciation and is treated as a textbook example of adaptive radiation, the rapid differentiation of a single ancestor into an . For example, do genes that evolve under positive selection in adaptive radiations of African cichlids also contribute to the speciation of Neotropical cichlids, or vice versa? The Poor man’s tropheus [Hypsophrys (Neetroplus) nematopus], which scrapes Aufwuchs from rocks, has an outer row of spatulate, incisor-like teeth with one to three inner rows of similarly shaped teeth. Speciation in cichlid fishes involved extremely high levels of morphological divergence, primarily related to ecologically relevant variation in body shape and size, and the trophic apparatus [13]. Repeated evolution of the same phenotypic difference during independent episodes of speciation is strong evidence for selection during speciation. Finally, it should be mentioned that polymorphism can be discussed only with respect to alleles of the same gene and not alleles of different genes at different loci. Zaki et al. From the sheer size of the family to the complexity of their ecological interactions and rapid evolution, cichlids provide a unique glimpse of the many factors that promote speciation. Although no difference was found in expression levels of the xanthophore-related candidate gene colony-stimulating factor 1 receptor a (csf1ra) between yellow and blue skin of P. saulosi [40], csf1ra is involved in the yellow pigmentation of the egg dummy colour patterning in other African cichlids [41]. The Tanganyika cichlid species flock thus provides an excellent model system to elucidate the evolutionary mechanisms which induce and trigger explosive speciation events. For this reason, the Neotropical adaptive radiation of Midas cichlid fishes (A. citrinellus species complex) is an ideal geographical, ecological and biological system in which to study the genomics of adaptation and speciation (reviewed in [15]). Since this trait is commonly differentiated - among other traits - between Midas Cichlid species, its plasticity might be an important factor in Midas Cichlid speciation. Abdel-Fattah M. El-Sayed, in Tilapia Culture (Second Edition), 2020. (a) A breeding pair of Midas cichlids (Amphilophus xiloaensis) from crater Lake Xiloá, Nicaragua. The models are based on a combination of laboratory experiments and field observations on the ecology, male and female mating behavior, and inheritance . Pseudomonas spp. Found inside – Page 389Speciation . Pp . 86–102 in Cichlid fishes : behaviour , ecology and evolution , M. H. Keenleyside , ed . London , England : Chapman and ... Similar morphologies of cichlid fish in Lakes Tanganyika and Malawi are due to convergence . Our study includes phylogeographic, population-genetic (based on mitochondrial DNA, microsatellites and amplified . Ten fish species have been reported and the four most abundant are the tui chub (Gila bicolor), Tahoe sucker (Catostomus tahoensis), Lahontan cutthroat trout (Oncorhynchus clarki henshawi), and cui-ui (Chasmistes cujus), which are endemic to these lakes (Galat et al., 1981). Mol Phylogenet Evol. Contains selected papers presented at a symposium at Jinja, Uganda in February 1993 which was organized by IDEAL, the International Decade for East African Lakes, a ten-year, multi- national, multi-disciplinary investigation of the ... Table 1.1. Thanks to M. Pierotti and three anonymous reviewers for comments improving the manuscript, and P. Eriksson, S. Balshine and A. Konings for contributing images. Modes and mechanisms of speciation in cichlid adaptive radiations The evidence for monophyly of lacustrine cichlid species flocks suggests that the vast majority of speciation events must have However, there is an increasing public awareness regarding the use of antimicrobials for disease control due to their human health and environmental problems. To understand sympatric speciation, one must first understand the other types of speciation. 1.9), jumping two positions since 2002. [37] found that, although the expression of the gene Mc1r (a common candidate gene for coloration [35]) was upregulated in the skin of gold fishes, comparative genomic analyses identified no sequence polymorphism in Mc1r between gold and dark Midas cichlids. Gunter et al. See §1b for more details. The Victoria and Malawi cichlids are all, without exception, maternal mouthbrooders (females brood their eggs by buccal incubation; reviewed by Barlow, 1991; Keenleyside, 1991), and the Tanganyika flock contains several lineages of mouthbrooders as well as substrate breeders (Nishida, 1991; Sturmbauer et al., 1994; Kocher et al., 1995). Basophils and eosinophils increased significantly in vitamin C–supplemented diets only (10%, 60%). nuptial coloration) have been shown to be a target of sexual selection by female mate choice both at the intra- and interspecific level , and female choice for male coloration has been suggested to promote speciation in African cichlids [20-23]. Thus, functional tests of these genes or more rigorous statistical methods based on population samples [88] are needed in future studies. Divergent Selection on Opsins Drives Incipient Speciation in Lake Victoria Cichlids. Putatively orthologous genes between species were determined by the bi-directional blast hit method. Soto-Rodriguez et al. The pharyngeal jaw apparatus of Midas Cichlids can be expressed plastically if stimulated mechanically during feeding. The skewness of tree length distributions as a measure of information content (Hillis and Huelsenbeck, 1992) was tested by generating 10,000 random trees. Cichlid. Blue male cichlid fish from a population that lack red phenotypes biased aggression towards blue stimulus male cichlid fish. Over the course of the past 15,000, an estimated over 500 different species of cichlid have evolved in Lake Victoria--equivalent to a new species developing every 30 years (Berkeley, 2009; Spinney, 2010). Most species have a row of unicuspid teeth on the premaxilla and dentary, with one to seven inner rows of smaller, unicusid teeth. DIJKSTRA, PETER., & Dijkstra, P. (2006). We observe that the oldest lineages of a species flock of cichlids are often less species-rich and live in the open water or deepwater habitats. Challenge tests revealed that P. anguilliseptica was the most pathogenic, inducing 97% mortality. The enormous number of species (1000 in Lake Malawi alone), the great diversity of trophic adaptations and behaviors, and the extreme rapidity of their divergence (<50,000 y for some faunas) single out these organisms as examples of evolution in progress . Cichlid fish from the tiny volcanic crate. Enter your email address below and we will send you your username, If the address matches an existing account you will receive an email with instructions to retrieve your username. Here, we review advances in the genomics and transcriptomics of cichlids, particularly regarding ecologically relevant differences in body shape, trophic apparatus, coloration and patterning, and sex determination. All comparisons gain tremendously in power if made in an explicit phylogenetic framework. 2015 Feb;83:56-71. doi: 10.1016/j.ympev.2014.10.009. Oral immunostimulant containing vitamin C could result in activating the immune system in a nonspecific way, thus providing resistance against pathogens. Many of the more than 2000 species have diversified based on ecological niche within lakes and often in parallel within and across radiating lineages [2]. Managua differently use the limnetic and . These tiny lakes may hold the key to understanding cichlid speciation, but they are extremely fragile habitats. The rapid radiations of extremely colorful cichlid fish in African lakes have fueled the hypothesis that sexual selection can drive species divergence without geographical isolation. Sequence analyses of mitochondrial DNA (mtDNA) (Meyer et al., 1990; Sturmbauer and Meyer, 1992, 1993; Kocher et al., 1993, 1995; Sturmbauer et al., 1994; Moran and Kornfield, 1993; Schliewen et al., 1994; Bowers et al., 1994) have extended the phylogenetic trees of the Lake Tanganyika and Malawi lineages and confirmed the monophyly of the Lake Victoria species flock. Martin CH, Feinstein LC. Lake Apoyo is a larger and older crater lake to the south. By using a microarray approach, it was shown that dominant and subordinate males differ significantly in their expression levels of almost 5 per cent of the tested genes, including co-regulated gene sets of neuroendocrine pathways [56]. Found inside – Page 357Mol Ecol 17:4500 Kocher TD (2004) Adaptive evolution and explosive speciation: the cichlid fish model. Nat Rev Genet 5:288–298 Kornfield I (1978) Evidence for rapid speciation in Africa cichlid fishes. Experientia 34:335–336 Kornfield I ... Would you like email updates of new search results? Different Sources of Allelic Variation Drove Repeated Color Pattern Divergence in Cichlid Fishes. Devashish Kar, in Epizootic Ulcerative Fish Disease Syndrome, 2016. Here, Winkelmann et al.show that competition between cichlid fish in Lake Tanganyika promotes . The influx of the Ruzizi River also ended a long period of isolation from the Zaire River system and caused an overflow of Lake Tanganyika via the Lukuga into the Lualaba, the upper reaches of the Zaire River.
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