Molecular phylogeny of the clariid fishes of Lake Victoria,Tanzania, inferred from cytochrome b DNA sequences

Mitochondrial DNA cytochrome b (cyt b) sequence variation among the clariid fishes of Lake Victoria and the Malagarasi wetland Tanzania were studied between August 2003 and February 2005. Seven species were sampled and together with 26 cyt b sequences from GenBank were used to reconstruct phylogenetic relationships in the family Clariidae. The study revealed two clades: one consisting of the big‐head species, Clarias gariepinus and Heterobranchus longifilis, and the other of small‐sized species, Clarias werneri, Clarias alluaudi, Clarias liocephalus and Clariallabes petricola. The study further revealed that the genus Clarias is paraphyletic and that H. longifilis clusters deeply inside the Clarias group. Although H. longifilis is thought to be the oldest in evolutionary age in Tanzania, it is not close to the ancestor of the Clariidae family. The results also showed that C. werneri and C. alluaudi are genetically distinct from each other.     

Gene flow between species of Lake Victoria haplochromine fishes

The haplochromine cichlid fishes of Lake Victoria (LV), East Africa, are a textbook example of adaptive radiation-a rapid divergence of multiple morphologically distinguishable forms from a few founding lineages. The forms are generally believed to constitute a "flock" of several hundred reproductively isolated species in a dozen or so genera. This belief has, until now, not been subjected to a test, however. Here, we compare genetic variation at 11 loci in 10 haplochromine populations of 6 different species. Although the genetic diversity in the populations is quite high, using a variety of statistical tests, we find no evidence of genetic differentiation among the populations of LV haplochromines. On genetic distance trees, populations of the same species intermingle with those of different species. At the molecular level, the species are indistinguishable from one another. Genetic comparisons with closely related species in 2 crater lakes indicate that the species within LV continue exchanging genes. These observations have important implications for phylogenetic reconstruction. The approach used in this study is applicable to other instances of adaptive radiation.     

High prevalence of non-synonymous substitutions in mtDNA of cichlid fishes from Lake Victoria

When a population size is reduced, genetic drift may fix slightly deleterious mutations, and an increase in nonsynonymous substitution is expected. It has been suggested that past aridity has seriously affected and decreased the populations of cichlid fishes in Lake Victoria, while geographical studies have shown that the water levels in Lake Tanganyika and Lake Malawi have remained fairly constant. The comparably stable environments in the latter two lakes might have kept the populations of cichlid fishes large enough to remove slightly deleterious mutations. The difference in the stability of cichlid fish population sizes between Lake Victoria and the Lakes Tanganyika and Malawi is expected to have caused differences in the nonsynonymous/synonymous ratio, ω (= d_N/d_S), of the evolutionary rate. Here, we estimated ω and compared it between the cichlids of the three lakes for 13 mitochondrial protein-coding genes using maximum likelihood methods. We found that the lineages of the cichlids in Lake Victoria had a significantly higher ω for several mitochondrial loci. Moreover, positive selection was indicated for several codons in the mtDNA of the Lake Victoria cichlid lineage. Our results indicate that both adaptive and slightly deleterious molecular evolution has taken place in the Lake Victoria cichlids' mtDNA genes, whose nonsynonymous sites are generally conserved.     

The conservation of the fishes of Lake Victoria,Africa: an ecological perspective

Synopsis The African Great Lakes are considered to be dynamically fragile ecosystems that are relatively resistant to minor changes with which they have co-evolved but vulnerable to major perturbations such as overfishing, the introduction of alien species and pollution. These lakes are inhabited by large species flocks of cichlid fishes which are characterised by a complex structure of interaction both between and within species, as is typical of mature ecosystems. Major perturbations, such as the disruption of trophic interactions through the introduction of alien fishes, may reverse the domination of relatively precocial, specialised forms and result in the creation of conditions that are conducive to the survival of more altricial, generalised forms with strong colonising abilities. The introduction of Nile perch and Nile tilapia, as well as other alien fishes, into Lake Victoria, combined with overfishing for the indigenous cichlid species, has resulted in marked changes to the fish communities and the fisheries that depend on them. The most important impacts of the Nile perch appear to be predation and aggressive effects whereas those of the tilapias include hybridization, overcrowding, competition for food and possibly the introduction of parasites and diseases. While the three proposed methods of conserving the indigenous flocks of cichlid fishes (captive propagation, reducing Nile perch stocks and closure of the haplochromine trawl fishery) all have merit, the changes that are occurring in Lake Victoria are basically irreversible. The highest priority should be to assist the governments of the riparian countries (Tanzania, Uganda and Kenya) with monitoring and research programmes and to support their policies of non-introduction of further alien fishes into any of the African Great Lakes so as prevent the same cycle of events from occurring, for example, in Lakes Tanganyika and Malawi. The diverse animal and plant communities of the African Great Lakes are a heritage of all mankind and it is the duty of every country to play a role in their conservation. It is therefore proposed that an internationally funded research programme should be mounted on the African Great Lakes on the scale of the tropical forest biome project of the IUCN. Editorial     

Microcystin production by cyanobacteria in the Mwanza Gulf (Lake Victoria,Tanzania)

In order to investigate the potential for microcystin (MC) production by cyanobacteria in the Mwanza Gulf (Lake Victoria, Tanzania), nutrients, phytoplankton and microcystins were sampled inshore (3 m depth) and offshore (18 m depth) from May to August 2002. Significant differences in soluble reactive phosphorus (SRP) and nitrate concentrations between offshore and inshore indicated eutrophication via terrestrial run-off. Though the concentrations of SRP and nitrate ranged between 36–127 and 35–726 μg l ⁻¹ each, the phytoplankton biovolume was generally low. The phytoplankton community was dominated by diatoms (Nitzschia acicularis), a number of cyanobacterial species (Aphanocapsa sp., Anabaena sp., Planktolyngbya spp., Microcystis sp.) and cryptomonads. The water column was completely mixed and Nitzschiapeaked in abundance during July. All cyanobacteria were low in abundance during the entire study period (0.1–1.6 mm ³ l ⁻¹). Microcystins were analysed using high performance liquid chromatography coupled with diode array detection High Performance Liquid Chromatography with Diode Array Detection (HPLC-DAD) and in most samples no microcystins were detected. The highest concentration of [Asp ³]-MC-RR was found in open water at the surface on July 2nd, 2002 (1 μg l ⁻¹). MC concentrations did not pose a potential health risk in the Mwanza Gulf during the study period, however, it is possible that the period of higher cyanobacterial biovolumes has been missed during the sampling period of this study.     

The origin and age of haplochromine fishes in Lake Victoria, east Africa

According to a widely held view, the more than 300 species of haplochromine cichlid fishes in Lake Victoria (LV), East Africa, originated from a single founder species in less than 12,000 years. This view, however, does not follow from the published geological and molecular evidence. The former does indeed suggest that the LV basin dried out less than 15,000 years ago, but it does not provide any information about the species that re-colonized the new lake or that remained in the rivers draining the area. The molecular evidence is inconclusive with respect to the origin of the LV haplochromines because cichlids from critical regions around LV were not adequately sampled; and as far as the age of the LV haplochromines is concerned, it in fact led to an estimate of 250,000-750,000 years old. In the present study, mitochondrial DNA (control region) variation was determined by heteroduplex and sequencing analyses of more than 670 specimens collected at widely distributed East African riverine and lacustrine localities. The analyses revealed the existence of seven haplogroups (I-VII) distinguishable by characteristic substitutions. All endemic LV samples tested fell into one of these haplogroups (V) which, however, was also found to be present at various other localities, both riverine and lacustrine, outside LV. Within this haplogroup, four subgroups (VA through VD) could be distinguished, two of which (VB and VC) were represented in LV and at other localities. The great majority of the LV haplochromine species could be classified as belonging to the VC subgroup, which was found only in LV and in the rivers draining into it. Hence, while the endemic haplochromine species of LV could not have originated from a single founding population, the lake does harbour a large species flock which probably arose in situ.     

Divergent parasite infections in sympatric cichlid species in Lake Victoria

Parasitism has been proposed as a factor in host speciation, as an agent affecting coexistence of host species in species‐rich communities and as a driver of post‐speciation diversification. Young adaptive radiations of closely related host species of varying ecological and genomic differentiation provide interesting opportunities to explore interactions between patterns of parasitism, divergence and coexistence of sympatric host species. Here, we explored patterns in ectoparasitism in a community of 16 fully sympatric cichlid species at Makobe Island in Lake Victoria, a model system of vertebrate adaptive radiation. We asked whether host niche, host abundance or host genetic differentiation explains variation in infection patterns. We found significant differences in infections, the magnitude of which was weakly correlated with the extent of genomic divergence between the host species, but more strongly with the main ecological gradient, water depth. These effects were most evident with infections of Cichlidogyrus monogeneans, whereas the only host species with a strictly crevice‐dwelling niche, Pundamilia pundamilia, deviated from the general negative relationship between depth and parasitism. In accordance with the Janzen–Connell hypothesis, we also found that host abundance tended to be positively associated with infections in some parasite taxa. Data on the Pundamilia sister species pairs from three other islands with variable degrees of habitat (crevice) specialization suggested that the lower parasite abundance of P. pundamilia at Makobe could result from both habitat specialization and the evolution of specific resistance. Our results support influences of host genetic differentiation and host ecology in determining infections in this diverse community of sympatric cichlid species.     

Prevalence of Schistosomes and Soil-Transmitted Helminths among Schoolchildren in Lake Victoria Basin,Tanzania

The objectives of this study was to conduct a survey on schistosomiasis and soil-transmitted helminth (STH) infections in order to come up with feasible control strategies in Lake Victoria basin, Tanzania. Depending on the size of the school, 150-200 schoolchildren were recruited for the study. Duplicate Kato-Katz stool smears were prepared from each child and microscopically examined for Schistosoma mansoni and STHs. Urine specimens were examined for Schistosoma haematobium eggs using the filtration technique. After the survey, mass drug administration was done using praziquantel and albendazole for schistosomiasis and STHs infections, respectively. A total of 5,952 schoolchildren from 36 schools were recruited for the study and had their stool and urine specimens examined. Out of 5,952 schoolchildren, 898 (15.1%) were positive for S. mansoni, 754 (12.6%) for hookworms, 188 (3.2%) for Ascaris lumblicoides, and 5 (0.008%) for Trichuris trichiura. Out of 5,826 schoolchildren who provided urine samples, 519 (8.9%) were positive for S. haematobium eggs. The results revealed that intestinal schistosomiasis, urogenital schistosomiasis, and STH infections are highly prevalent throughought the lake basin. The high prevalence of intestinal and urogenital schistosomisiasis in the study area was a function of the distance from Lake Victoria, the former being more prevalent at localities close to the lake, whilst the latter is more so away from it. Control of schistosomiasis and STHs in the study area requires an integrated strategy that involves provision of health education to communities, regular treatments, and provision of adequate safe water supply and sanitation facilities.     

The benthic macrofauna community of Kenyan waters of Lake Victoria

The benthic macroinvertebrates in the Kenya waters of Lake Victoria (ca. 1400 km²) were surveyed during four 10 day sampling periods in February, April, August and November 1984. Fourty three taxonomic groups were recorded. Oligochaetes, molluscs and dipteran larvae were the most abundant and widely distributed groups. The dominant oligochaetes were Branchiura sowerbyii Beddard and the swamp worm Alma emini Michaelsen, which were abundant in all silty and soft mud areas in both littoral and open water zones. Melanoides tuberculata Müller, Bellamya unicolor Olivier and Caelatura spp. were the most abundant molluscs. Different patterns were observed between near-littoral stations (<8 m deep) and deep water stations (8–40 m deep). After 10 years (1994), the macroinvertebrates of Lake Victoria were examined again. No evidence was found of seasonal changes in the composition or benthos density, although local changes occurred in some species which were attributed to swarming behaviour or larval settlement patterns. The spatial distribution of the fauna may be influenced primarily by oxygen availability and industrial effluent discharged from paper mill and agro-based industries in the catchment of the lake. There has been a shift in abundance, from an Oligochaeta and Insecta dominated community in 1984, to the present community dominated by Mollusca and Oligochaeta. The ecological role of the benthic community and its recent changes to fisheries production in the lake is discussed.     

Microhabitat distributions and species interactions of ectoparasites on the gills of cichlid fish in Lake Victoria,Tanzania

Heterogeneous exposure to parasites may contribute to host species differentiation. Hosts often harbour multiple parasite species which may interact and thus modify each other’s effects on host fitness. Antagonistic or synergistic interactions between parasites may be detectable as niche segregation within hosts. Consequently, the within-host distribution of different parasite taxa may constitute an important axis of infection variation among host populations and species. We investigated the microhabitat distributions and species interactions of gill parasites (four genera) infecting 14 sympatric cichlid species in Lake Victoria, Tanzania. We found that the two most abundant ectoparasite genera (the monogenean Cichlidogyrus spp. and the copepod Lamproglena monodi) were non-randomly distributed across the host gills and their spatial distribution differed between host species. This may indicate microhabitat selection by the parasites and cryptic differences in the host–parasite interaction among host species. Relationships among ectoparasite genera were synergistic: the abundances of Cichlidogyrus spp. and the copepods L. monodi and Ergasilus lamellifer tended to be positively correlated. In contrast, relationships among morphospecies of Cichlidogyrus were antagonistic: the abundances of morphospecies were negatively correlated. Together with niche overlap, this suggests competition among morphospecies of Cichlidogyrus. We also assessed the reproductive activity of the copepod species (the proportion of individuals carrying egg clutches), as it may be affected by the presence of other parasites and provide another indicator of the species specificity of the host–parasite relationship. Copepod reproductive activity did not differ between host species and was not associated with the presence or abundance of other parasites, suggesting that these are generalist parasites, thriving in all cichlid species examined from Lake Victoria.     

Organochlorine residues in fish from Lake Victoria, Kenya

Eighty-two samples of either nile perch fish fat or muscle were collected from the Kenyan region of Lake Victoria for detection of organochlorine residues. Nine organochlorine residues were detected in the following percentages: α-BHCIHCB-40%; P-BHC/HCB-40%; y-BHC/HCB/lindane-4%; aldrin-9%; dieldrin-1%; p, p'-DDE-73%; p, p'-DDD-9%; o, p'-DDT-170; and p, p'-DDT-11%. All levels of organochlorine residues were below the Extraneous Residue Limit (ERL), apart from just one sample of fish fat which had 4.51 ppm of sum DDT above ERL. There was a higher level of organochlorines from Mbita than from Luanda fishing point.

Résumé

Eighty-two échantillons de graisse ou de muscle de perche du Nil ont été récoltés dans la région kényane du lac Victoria pour rechercher des résidus organochlorés. On a détecté 9 résidus organochlorés, avec les pourcentages suivants: α-BHC/HCB-40%; β-BHC/HCB-40%;γ-BHC/HCB/lindane-4%;aldrine-9%; dieldrine-1%; p, p'-DDE-73%; p, p'-DDD-9%; o, p'-DDT-1%; and p, p'-DDT-11%. Tous les taux de résidus organochlorés ktaient infkrieurs a la 'Extraneous Residue Limit' (ERL) sauf pour un seul échantillon de graisse de poisson qui contenait 4,51 ppm de DDT total au-dessus de 1'ERL. I1 y avait un taux plus élevé d'organochlorés a Mbita qu'au point de pêche de Luanda.     

On the age and origin of the species flock of haplochromine cichlid fishes of Lake Victoria

Recent suggestions concerning the age and origin of the flock of haplochromine cichlid fishes in Lake Victoria (East Africa) are considered. These accept as proven the suggestion that Lake Victoria dried out completely in the Late Pleistocene, was dry for several thousand years, and refilled ca. 12400 years ago. Apart from the fact that other geophysical evidence contradicts this claim, its biological implications, which do likewise, have never been considered by those who have accepted it. Like those of all previous authors who have seized on the presence of the haplochromine flock of perhaps more than 600 species as evidence of extremely rapid evolution since the lake allegedly refilled, the account completely overlooks the fact that any such desiccation must have eliminated not only the haplochromine cichlids but the entire biota of the lake. Nevertheless, its present fauna not only includes the haplochromines but many other endemic organisms that would not be expected, and whose presence and history demand an explanation if the lake did indeed dry out. No such explanation has been offered, nor does such seem possible. The recent interpretation of events is questioned and rejected.     

Phytoplankton community structure and environment in the Kenyan waters of Lake Victoria

• 1 Phytoplankton species composition, numerical abundance, spatial distribution and total biomass measured as chlorophyll a concentration were studied in relation to environmental factors in September 1994 (dry season) and March 1995 (rainy season), respectively, in the Kenyan waters of Lake Victoria; 103 species were recorded.
• 2 Blue‐green algae (Cyanophyceae) were most diverse, followed by diatoms (Bacillariophyceae), green algae (Chlorophyceae) and dinoflagellates (Dinophyceae).
• 3 Twinspan separated the phytoplankton communities in the Nyanza Gulf and those in the open lake during both seasons. During the dry season, the Nyanza Gulf was strongly dominated by blue‐greens, while diatoms dominated in the open lake. During the rainy season, blue‐greens remained dominant in the Nyanza Gulf although the number of species found was lower than during the dry season; in the open lake, blue‐greens replaced diatoms as the dominant group and there were more species than in the dry season.
• 4 Canonical correspondence analysis indicated that the phytoplankton species distribution was significantly correlated with turbidity during the dry season and with SiO₂ during the rainy season. Chlorophyll a concentrations ranging from 2.0 to 71.5 mg m^‐3 in the dry season and 2.0–17.2 mg m^‐3 in the rainy season confirm earlier reports of increasing phytoplankton biomass in Lake Victoria since the 1960s.

     

Downstream changes in the composition of the parasite community of fishes in an Appalachian stream

The spatial distribution of 6 parasite species (Myxobolus sp., Dactylogyrus sp., Sterliadochona ephemeridarum, Plagioporus sinitsini, Allopodocotyle chiliticorum, Allocreadium lucyae) was studied in 5 species of fishes (Oncorhynchus mykiss, Clinostomus funduloides, Notropis chiliticus, Rhinichthys atratulus, Semotilus atromaculatus) in Basin Creek, an Appalachian stream in North Carolina. Nonmetric multidimensional scaling and vector fitting were used to determine if the proximity of sampling sites was related to community similarity. Position along Basin Creek was significantly related to parasite community structure. Breaks in parasite community composition were imposed by waterfalls at upstream areas of Basin Creek that restricted distributions of C. funduloides, N. chiliticus, and S. atromaculatus and at the downstream limit of the study area by a break in the distribution of S. ephemeridarum coincident with the existence of a dam but were independent of suitable piscine host distributions. These discontinuities in parasite community composition imply that the relationship between proximity of sites and community similarity is predictive because distance between sites is related to the probability that fish at different sampling sites recruit parasites from different species pools. This relationship is not the same for all component communities.     

Male-male competition and nuptial-colour displacement as a diversifying force in Lake Victoria cichlid fishes

We propose a new mechanism for diversification of male nuptial-colour patterns in the rapidly speciating cichlid fishes of Lake Victoria. Sympatric closely related species often display nuptial colours at opposite ends of the spectrum with males either blue or yellow to red. Colour polymorphisms within single populations are common too. We propose that competition between males for breeding sites promotes such colour diversification, and thereby speciation. We hypothesize that male aggression is primarily directed towards males of the common colour, and that rare colour morphs enjoy a negatively frequency-dependent fitness advantage. We test our hypothesis with a large dataset on the distributions and nuptial colorations of 52 species on 47 habitat islands in Lake Victoria, and with a smaller dataset on the within-spawning-site distributions of males with different coloration. We report that territories of males of the same colour are negatively associated on the spawning site, and that the distribution of closely related species over habitat islands is determined by nuptial coloration in the fashion predicted by our hypothesis. Whereas among unrelated species those with similar nuptial colour are positively associated, among closely related species those with similar colour are negatively associated and those with different colour are positively associated. This implies that negatively frequency-dependent selection on nuptial coloration among closely related species is a sufficiently strong force to override other effects on species distributions. We suggest that male-male competition is an important and previously neglected agent of diversification among haplochromine cichlid fishes.     

Species inventory and the local uses of the plants and fishes of the Lower Sondu Miriu wetland of Lake Victoria,Kenya

The release of phosphate from anoxic sediments is still not precisely understood. The proposal by Einsele (1936), later adapted by Mortimer (1941, 1942), that this release is caused by the reduction of a FeOOH-phosphate complex, is generally accepted as the reaction mechanism, although there is no experimental evidence for it. Golterman (1995a) and De Groot (1991) have shown that this P-release may indeed be brought about by H₂S, but only if a large excess of H₂S is available. In lakes, however, the reducing capacity is relatively small as most of the organic carbon produced by primary production is used for other reduction processes. The solubilization of apatite is a likely alternative, as anoxic conditions are automatically concomitant with a pH decrease, and in hard waters the formation of apatite is well demonstrated. This mechanism is not active in soft waters, such as those studied by Mortimer. Another proposed process is the release of polyphosphate by sediment bacteria. Experimental evidence for this mechanism is, however, weak. The fourth possibility is the need for bacteria to mineralize a larger part of the sediment organic matter under anoxic conditions in order to obtain the same amount of energy, as some energy will be retained in the fermentation products. There is circumstantial evidence for this hypothesis, but laboratory experiments are needed before real evidence will be available.     

Characterization of Cryptocaryon irritans,a parasite isolated from marine fishes in Taiwan

The ciliated protozoan parasite Cryptocaryon irritans infecting marine fishes in Taiwan is described. Developmental characteristics and sequences of the ribosomal DNA regions such as part of 18 S, the entire first internal transcribed spacer, and part of 5.8 S of various Taiwan isolates of C. irritans were investigated. A total of 5 isolates was obtained from different fish-host species and localities, the majority from cultured fish species. C. irritans from Taiwan is able to shift its developmental characteristics, i.e. from non-adherent to adherent tomonts, from individualistic to aggregate-forming tomonts, from infection of the gills only to infection of the gills and body. Thus, it is not possible to classify strains of C. irritans on the basis of these parameters. Premature tomonts that developed from dead fishes were able to produce theronts that could infect fish host. Isolates from Pingtung and the USA had identical nucleotide sequences while an isolate from Malaysia was identical to an Israel isolate. Percentage variation among pairs of Taiwan isolates showed a higher degree of variation than isolate sequences listed in GenBank. Sequence analysis revealed highly aberrant isolates in Taiwan, and a phylogenetic tree distinguished a marine and a low-salinity variant. C. irritans from marine fishes in Taiwan, therefore, display some characteristics not previously reported. Since manipulation of salinity in brackishwater ponds and marine cage sites is not feasible, there is a need to develop new strategies for the control and prevention of cryptocaryoniasis.     

Dynamics and limitations of phytoplankton biomass along a gradient in Mwanza Gulf,southern Lake Victoria (Tanzania)

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B chromosomes have a functional effect on female sex determination in Lake Victoria cichlid fishes

The endemic cichlid fishes in Lake Victoria are a model system for speciation through adaptive radiation. Although the evolution of the sex-determination system may also play a role in speciation, little is known about the sex-determination system of Lake Victoria cichlids. To understand the evolution of the sex-determination system in these fish, we performed cytogenetic analysis in 11 cichlid species from Lake Victoria. B chromosomes, which are present in addition to standard chromosomes, were found at a high prevalence rate (85%) in these cichlids. In one species, B chromosomes were female-specific. Cross-breeding using females with and without the B chromosomes demonstrated that the presence of the B chromosomes leads to a female-biased sex ratio in this species. Although B chromosomes were believed to be selfish genetic elements with little effect on phenotype and to lack protein-coding genes, the present study provides evidence that B chromosomes have a functional effect on female sex determination. FISH analysis using a BAC clone containing B chromosome DNA suggested that the B chromosomes are derived from sex chromosomes. Determination of the nucleotide sequences of this clone (104.5 kb) revealed the presence of several protein-coding genes in the B chromosome, suggesting that B chromosomes have the potential to contain functional genes. Because some sex chromosomes in amphibians and arthropods are thought to be derived from B chromosomes, the B chromosomes in Lake Victoria cichlids may represent an evolutionary transition toward the generation of sex chromosomes.