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PubMed2005

Parallelism of amino acid changes at the RH1 affecting spectral sensitivity among deep-water cichlids from Lakes Tanganyika and Malawi.

Many examples of the appearance of similar traits in different lineages are known during the evolution of organisms. However, the underlying genetic mechanisms have been elucidated in very few cases. Here, we provide a clear example of evolutionary parallelism, involving changes in the same genetic pathway, providing functional adaptation of RH1 pigments to deep-water habitats during the adaptive radiation of East African cichlid fishes. We determined the RH1 sequences from 233 individual cichlids. The reconstruction of cichlid RH1 pigments with 11-cis-retinal from 28 sequences showed that the absorption spectra of the pigments of nine species were shifted toward blue, tuned by two particular amino acid replacements. These blue-shifted RH1 pigments might have evolved as adaptations to the deep-water photic environment. Phylogenetic evidence indicates that one of the replacements, A292S, has evolved several times independently, inducing similar functional change. The parallel evolution of the same mutation at the same amino acid position suggests that the number of genetic changes underlying the appearance of similar traits in cichlid diversification may be fewer than previously expected.

Adaptation, Physiological↗

PubMed2001

Lake level fluctuations synchronize genetic divergences of cichlid fishes in African lakes.

Water level fluctuations are important modulators of speciation processes in tropical lakes, in that they temporarily form or break down barriers to gene flow among adjacent populations and/or incipient species. Time estimates of the most recent major lowstands of the three African Great Lakes are thus crucial to infer the relative timescales of explosive speciation events in cichlid species flocks. Our approach combines geological evidence with genetic divergence data of cichlid fishes from the three Great East African Lakes derived from the fastest-evolving mtDNA segment. Thereby, we show for each of the three lakes that individuals sampled from several populations which are currently isolated by long geographic distances and/or deep water form clusters of equally closely related haplotypes. The distribution of identical or equally closely related haplotypes in a lake basin allows delineation of the extent of lake level fluctuations. Our data suggest that the same climatic phenomenon synchronized the onset of genetic divergence of lineages in all three species flocks, such that their most recent evolutionary history seems to be linked to the same external modulators of adaptive radiation. A calibration of the molecular clock of the control region was elaborated by gauging the age of the Lake Malawi species flock through the divergence among the utaka-cichlid and the mbuna-cichlid lineages to minimally 570,000 years and maximally 1 Myr. This suggests that the low-lake-level period which established the observed patterns of genetic relatedness dates back less than 57,000 years, probably even to 17,000-12,400 years ago, when Lake Victoria dried up and Lakes Malawi and Tanganyika were also low. A rapid rise of all three lakes about 11,000 years ago established the large-scale population subdivisions observed today. Over that period of time, a multitude of species originated in Lakes Malawi and Victoria with an impressive degree of morphological and ecological differentiation, whereas the Tanganyikan taxa that were exposed to the same habitat changes hardly diverged ecologically and morphologically. Our findings also show that patterns of genetic divergences of stenotopic organisms provide valuable feedback on geological and sedimentological time estimates for lake level changes.

Africa↗

PubMed2001

Cone opsin genes of african cichlid fishes: tuning spectral sensitivity by differential gene expression.

Spectral tuning of visual pigments is typically accomplished through changes in opsin amino acid sequence. Within a given opsin class, changes at a few key sites control wavelength specificity. To investigate known differences in the visual pigment spectral sensitivity of the Lake Malawi cichlids, Metriaclima zebra (368, 488, and 533 nm) and Dimidiochromis compressiceps (447, 536, and 569 nm), we sequenced cone opsin genes from these species as well as Labeotropheus fuelleborni and Oreochromis niloticus. These cichlids have five distinct classes of cone opsin genes, including two unique SWS-2 genes. Comparisons of the inferred amino acid sequences from the five cone opsin genes of M. zebra, D. compressiceps, and L. fuelleborni show the sequences to be nearly identical. Therefore, evolution of key opsin sites cannot explain the differences in visual pigment sensitivities. Real-time PCR demonstrates that different cichlid species express different subsets of the available cone opsin genes. Metriaclima zebra and L. fuelleborni express a complement of genes which give them UV-shifted visual pigments, while D. compressiceps expresses a different set to produce a red-shifted visual system. Thus, variations in cichlid spectral sensitivity have arisen through evolution of gene regulation, rather than through changes in opsin amino acid sequence.

Animals↗

PubMed2001

The oldest fossil cichlids (Teleostei: Perciformes): indication of a 45 million-year-old species flock.

Five closely related species of fossil cichlids collected from an Eocene site in Tanzania, East Africa, represent the oldest known cichlids. The specimens are whole-body, articulated fishes that are extremely well preserved and, therefore, have the potential to add to our knowledge of the history of this family. Modern cichlids are particularly well known for the numerous species flocks of the East African Great Lakes. A great deal of research is ongoing regarding all aspects of the fishes in these flocks, including their evolutionary history The new collection of fossils reported here is interpreted as representing a species flock that arose in a small crater lake. These fossils indicate that cichlids' ability to form species flocks evolved early in the history of this family.

Africa, Eastern↗

PubMed2001

Microsatellites reveal high levels of population substructuring in the species-poor Eretmodine cichlid lineage from Lake Tanganyika.

This study investigated fine-scale population substructuring in an apparently monogamous, biparental mouth-brooding cichlid. Microsatellite allele frequencies were determined at four polymorphic loci for nine populations of Eretmodus cyanostictus. We provide empirical support for the hypothesis that a species employing this breeding strategy should exhibit high levels of population substructuring. Stretches of sand represent considerable barriers to dispersal and, in contrast to the rock-dwelling cichlids of Lake Malawi, distance alone, along a continuous rocky shoreline, is sufficient to reduce gene flow significantly There was a significant pattern of isolation by distance both along the whole study area and over the stretch of continuous shoreline, suggesting that this species has poor dispersal capabilities and that juveniles establish territories close to their natal site. Despite limited dispersal, E. cyanostictus populations are not significantly more inbred than a more-widely dispersing rock-dwelling cichlid from Lake Malawi. This finding may cast doubt on the hypothesis that polyandry has evolved as a mechanism for maintaining genetic diversity in Lake Malawi cichlids. High levels of substructuring may not always promote high levels of speciation, and other factors, such as the intensity of sexual selection, may be more important in determining the speciation potential of a lineage.

Alleles↗

PubMedDate not supplied

[Feeding habits of cichlid species (Perciformes: Cichlidae) in Caobas lake, Quintana Roo, Mexico].

Feeding habits of seven cichlid species (Archocentrus octofasciatus, A. spilurus, "Cichlasoma" robertsoni, "C." synspilum, "C." urophthalmus, Petenia splendida, Thorichthys meeki) in Lake Caobas, southern Yucatan Peninsula, were studied. Samples were taken with enclosure and cast nets during the dry and rainy seasons of 1995 (day and night). The environment was characterized by measuring temperature, conductivity and pH. All individuals were below 41 mm SL (N = 281). Frequency of occurrence and prey abundance were analyzed. Main prey items were chironomids, mites, copepods, cladocerans, and ostracods. The cichlids fed mainly on zooplankton, with the partial exceptions of P. splendida (piscivore), "C." synspilum and A. spilurus (herbivores). A cluster analysis showed that the most similar trophic spectra were those of T. meeki, "C." robertsoni and "C." salvini, which were also the least diverse. "C." synspilum and A. spilurus had an intermediate distance between their diets and those of other species. The species with the most distinctive feeding composition were P. splendida (with the most diverse and equitable diet) and the omnivore A. octofasciatus (whose diet was the richest one). T. meeki showed quantitative diel, ontogenetic, and seasonal diet changes, but none between sexes. "C." robertsoni, "C." salvini and "C." synspilum differ in food habits in Caobas and in other localities, a fact that underscores the trophic adaptability of cichlids. Trophic overlap between cichlids in Caobas could imply absence of competition, perhaps because resources are abundant in the ecosystem.

Animals↗

PubMedDate not supplied

[Larval Clinostomum infection of cichlid fish in the lake of Kompienga in Burkina Faso: a menace to haleutic exploitation and public health].

Clinostomum sp. is a trematode, a parasitic worm, of which the larva infest a number of species of wild fish. In intertropical Africa and particularly in artificial lakes in the east of Burkina Faso, the infestation is especially observed in the cichlids fish. Because cichlids constitute 80 to 90% of the fish catches, enzootic infestation could infringe upon the fisheries, and also adversely affect public health. For this epidemiological study conducted during the first three quarters of 1994, we examined a total of 932 cichlids, a mix of all species. Prevalence (Pr) and intensity (I, number of parasites per fish) were the two indicators of the infestation level. 387 fish were carriers of the parasite, comprising 41.5% of the total. Differences were observed between the species of cichlids. Oreochromis niloticus was the most infested (Pr, 56%; I, 17), followed by Sarotherodon galilaeus (Pr, 44%; I,6); and Hemichromis fasciatus (Pr, 20%; I, 5). In 89% of the carriers, the parasite was localized on the internal face of the operculum and in the pericardial septum, compared to 11% of the localizations in subcutaneous and muscular tissues. These last two localizations render the fish repugnant, and can cause the fresh fish to be rejected. If the parasitized fish is not rejected and then insufficiently cooked, as occurs at public grills with multitudes of consumers, the living parasite can penetrate and stay in the upper digestive tract and constitute a threat for consumer health.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed2006

A BAC library of the East African haplochromine cichlid fish Astatotilapia burtoni.

A BAC library was constructed from Astatotilapia burtoni, a haplochromine cichlid that is found in Lake Tanganyika, East Africa, and its surrounding rivers. The library was generated from genomic DNA of blood cells and comprises 96,768 individual clones. Its median insert size is 150 kb and the coverage is expected to represent about 14 genome equivalents. The coverage evaluation was based on genome size estimates that were obtained by flow cytometry. In addition, hybridization screens with five probes largely corroborate the above coverage estimate, although the number of clones ranged from 5 to 22 authenticated clones per single copy probe. The BAC library described here is expected to be useful to the scientific community interested in cichlid genomics as an important resource to gain new insights into the rapid evolution of the great species diversity of haplochromine cichlid fishes.

Africa, Eastern↗

PubMed2006

Evolution of novelty in the cichlid dentition.

The shape of teeth occupies a central position in various biological disciplines, from paleo-ecology to molecular biology to cosmetic and reconstructive dentistry. Despite a long tradition of study in mammals, important questions remain regarding the genetic and developmental basis of differences in tooth shape. Here, we use natural mutants of cichlid fish from East Africa, which exhibit tremendous dental diversity, to help fill the gaps in our understanding of vertebrate odontogenesis. We employ an expanded genetic linkage map to demonstrate that cusp number segregates as a gene of major effect, which explains approximately 40% of the phenotypic variance, on cichlid chromosome 5. Furthermore, we examine patterns of Bmp4 expression in early odontogenesis to address and refine predictions of models linking tooth shape and tooth number. Mutations in the Bmp4 cistron do not control tooth shape in this mapping cross. Our data suggest that the evolution of novelty in the cichlid dentition is galvanized by a small number of genetic changes, echoing similar conclusions from recent studies of other vertebrate adaptive morphologies.

Animals↗

PubMed2003

Comparison of testes structure, spermatogenesis, and spermatocytogenesis in young, aging, and hybrid cichlid fish (Cichlidae, Teleostei).

Testis structure, spermatogenesis, and spermatocytogenesis were compared in 13 species of cichlid fishes, belonging to the subfamilies Haplochrominae and Tilapinae. The species studied were either mouth brooders, in which fertilization occurs mostly inside the mouth of the brooding fish, or substrate brooders, whose eggs adhere to a substrate over which the sperm is ejaculated. In this study, the embryogenesis of testes anlagen and sperm production was followed in embryos and in fish up to 15 years old, as well as in hybrids of the two subfamilies. In cichlids, the testes are of the unrestricted type and primary spermatogonia develop along the entire length of the developing sperm tubule. The first primary spermatogonia are observed in the testes anlagen 2-5 days after fertilization and they continue to develop in cysts formed by the enveloping Sertoli cells and the intertubular elements. The dimensions of such primary and secondary spermatocysts are correlated with the number of spermatogonia they contain and the corresponding number of mitotic multiplications. The largest mature cysts attained 300 microm, and contained 2,200-2,400 spermatids in the mouth-brooding species and 2,600-3,200 in the substrate-brooding species. Despite the fact that in such cysts cytoplasmic bridges connect only the isogamete spermatids, the maturation of all cells and consequent spermiation is synchronized. Meristic characters distinguish the sperm of mouth brooders from those of substrate brooders, especially in the number of mitochondria and length of the flagellum. In older fish and hybrids, various changes can be seen in the gametogenic epithelium and intertubular cells. These include thickening of the connective tissue, formation of "yellow" groups of Leydig cells, cell apoptosis and degeneration, and, especially, formation of large spermatogonia, with large, electron-dense nucleoli, that have the cytological characteristics of oocytes. The intra- and interspecific variability of sperm dimensions in the studied cichlids poses an interesting question in the context of sperm competition.

Aging↗

PubMed2006

Species-specific population structure in rock-specialized sympatric cichlid species in Lake Tanganyika, East Africa.

Species richness and geographical phenotypic variation in East African lacustrine cichlids are often correlated with ecological specializations and limited dispersal. This study compares mitochondrial and microsatellite genetic diversity and structure among three sympatric rock-dwelling cichlids of Lake Tanganyika, Eretmodus cyanostictus, Tropheus moorii, and Ophthalmotilapia ventralis. The species represent three endemic, phylogenetically distinct tribes (Eretmodini, Tropheini, and Ectodini), and display divergent ecomorphological and behavioral specialization. Sample locations span both continuous, rocky shoreline and a potential dispersal barrier in the form of a muddy bay. High genetic diversity and population differentiation were detected in T. moorii and E. cyanostictus, whereas much lower variation and structure were found in O. ventralis. In particular, while a 7-km-wide muddy bay curtails dispersal in all three species to a similar extent, gene flow along mostly continuous habitat appeared to be controlled by distance in E. cyanostictus, further restricted by site philopatry and/or minor habitat discontinuities in T. moorii, and unrestrained in O. ventralis. In contrast to the general pattern of high gene flow along continuous shorelines in rock-dwelling cichlids of Lake Malawi, our study identifies differences in population structure among stenotopic Lake Tanganyika species. The amount of genetic differentiation among populations was not related to the degree of geographical variation of body color, especially since more phenotypic variation is observed in O. ventralis than in the genetically highly structured E. cyanostictus.

Animals↗

PubMed2006

Time-place learning in individually reared angelfish, but not in pearl cichlid.

Time-place learning based on food association was investigated in the cichlids angelfish (Pterophyllum scalare) and pearl cichlid (Geophagus brasiliensis) reared in isolation, therefore eliminating social influence on foraging. During a 30-day period, food was placed in one side of the aquarium (containing three compartments) in the morning and in the opposite side in the afternoon. Learning was inferred by the number of correct side choices of all fish in each day of test (15th and 30th). During the test day fish were not fed. The angelfish learned to switch sides at the correct day period in order to get food, suggesting this species has time-place learning ability when individually reared. On the other hand, the same was not observed for pearl cichlid.

Animals↗

PubMed2002

Speciation via introgressive hybridization in East African cichlids?

Speciation caused by introgressive hybridization occurs frequently in plants but its importance remains controversial in animal evolution. Here we report a case of introgressive hybridization between two ancient and genetically distinct species of Lake Tanganyika cichlids that led to the formation of a new species. Neolamprologus marunguensis contains mtDNA haplotypes from both parental species varying on average by 12.4% in the first section of the control region and by 5.2% in a segment of the cytochrome b gene. All individuals have almost identical DNA sequences in the flanking regions of the single-copy nuclear DNA locus TmoM27, and show a mosaic of alleles derived from both parental lineages in six microsatellite loci. Hence, our finding displays another mode of speciation in cichlid fishes. The increase of genetic and phenotypic diversity due to hybridization may contribute to the uniquely rapid pace of speciation in cichlids.

Animals↗

PubMed2003

Genome mapping of the orange blotch colour pattern in cichlid fishes.

The dramatic variation of cichlid fish colour pattern is thought to function in mate choice, evolve by sexual selection, and contribute to explosive speciation. Here, we combine linkage mapping and population genetic analyses to identify a single region of the cichlid genome responsible for the orange blotch (OB) colour phenotype. In each analysis, OB is tightly linked to the c-ski1 gene. Additionally, we use comparative mapping information from the Takifugu rubripes and human genomes to suggest positional candidate loci for OB. Our work should engender a more comprehensive understanding of the molecular ecology of OB and its role in cichlid speciation. Moreover, we have assembled the components of a method to focus upon the genetic basis of evolutionarily and ecologically significant phenotypes.

Animals↗

PubMed2002

Patterns in fish radiation are compatible with Pleistocene desiccation of Lake Victoria and 14,600 year history for its cichlid species flock.

Geophysical data are currently being interpreted as evidence for a late Pleistocene desiccation of Lake Victoria and its refilling 14,600 years ago. This implies that between 500 and 1000 endemic cichlid fish species must have evolved in 14,600 years, the fastest large-scale species radiation known. A recent review concludes that biological evidence clearly rejects the postulated Pleistocene desiccation of the lake: a 14,600 year history would imply exceptionally high speciation rates across a range of unrelated fish taxa. To test this suggestion, I calculated speciation rates for all 41 phylogenetic lineages of fish in the lake. Except for one cichlid lineage, accepting a 14 600 year history does not require any speciation rates that fall outside the range observed in fishes in other young lakes around the world. The exceptional taxon is a lineage of haplochromine cichlids that is also known for its rapid speciation elsewhere. Moreover, since it is unknown how many founding species it has, it is not certain that its speciation rates are really outside the range observed in fishes in other young lakes. Fish speciation rates are generally faster in younger than in older lakes, and those in Lake Victoria, by far the largest of the young lakes of the world, are no exception. From the speciation rates and from biogeographical observations that Lake Victoria endemics, which lack close relatives within the lake basin, have such relatives in adjacent drainage systems that may have had Holocene connections to Lake Victoria, I conclude that the composition of the fish assemblage does not provide biological evidence against Pleistocene desiccation. It supports a hypothesis of recent colonization from outside the lake basin rather than survival of a diverse assemblage within the basin.

Africa, Eastern↗

PubMed2003

Nuclear markers reveal unexpected genetic variation and a Congolese-Nilotic origin of the Lake Victoria cichlid species flock.

Phylogenetic analyses based on mitochondrial (mt) DNA have indicated that the cichlid species flock of the Lake Victoria region is derived from a single ancestral species found in East African rivers, closely related to the ancestor of the Lake Malawi cichlid species flock. The Lake Victoria flock contains ten times less mtDNA variation than the Lake Malawi radiation, consistent with current estimates of the ages of the lakes. We present results of a phylogenetic investigation using nuclear (amplified fragment length polymorphism) markers and a wider coverage of riverine haplochromines. We demonstrate that the Lake Victoria-Edward flock is derived from the morphologically and ecologically diverse cichlid genus Thoracochromis from the Congo and Nile, rather than from the phenotypically conservative East African Astatotilapia. This implies that the ability to express much of the morphological diversity found in the species flock may by far pre-date the origin of the flock. Our data indicate that the nuclear diversity of the Lake Victoria-Edward species flock is similar to that of the Lake Malawi flock, indicating that the genetic diversity is considerably older than the 15 000 years that have passed since the lake began to refill. Most of this variation is manifested in trans-species polymorphisms, indicating very recent cladogenesis from a genetically very diverse founder stock. Our data do not confirm strict monophyly of either of the species flocks, but raise the possibility that these flocks have arisen from hybrid swarms.

Africa, Eastern↗

PubMed2006

Colour-assortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa.

The species flocks of cichlid fishes in the East African Lakes Tanganyika, Malawi and Victoria are prime examples of adaptive radiation and explosive speciation. Several hundreds of endemic species have evolved in each of the lakes over the past several thousands to a few millions years. Sexual selection via colour-assortative mating has often been proposed as a probable causal factor for initiating and maintaining reproductive isolation. Here, we report the consequences of human-mediated admixis among differentially coloured populations of the endemic cichlid fish Tropheus moorii from several localities that have accidentally been put in sympatry in a small harbour bay in the very south of Lake Tanganyika. We analysed the phenotypes (coloration) and genotypes (mitochondrial control region and five microsatellite loci) of almost 500 individuals, sampled over 3 consecutive years. Maximum-likelihood-based parenthood analyses and Bayesian inference of population structure revealed that significantly more juveniles are the product of within-colour-morph matings than could be expected under the assumption of random mating. Our results clearly indicate a marked degree of assortative mating with respect to the different colour morphs. Therefore, we postulate that sexual selection based on social interactions and female mate choice has played an important role in the formation and maintenance of the different colour morphs in Tropheus, and is probably common in other maternally mouthbrooding cichlids as well.

Alleles↗

PubMed2006

Axes of differentiation in the bower-building cichlids of Lake Malawi.

The 500-1000 cichlid species endemic to Lake Malawi constitute one of the most rapid and extensive radiations of vertebrates known. There is a growing debate over the role natural and sexual selection have played in creating this remarkable assemblage of species. Phylogenetic analysis of the Lake Malawi species flock has been confounded by the lack of appropriate morphological characters and an exceptional rate of speciation, which has allowed ancestral molecular polymorphisms to persist within species. To overcome this problem we used amplified fragment length polymorphism (AFLP) to reconstruct the evolution of species within three genera of Lake Malawi sand-dwelling cichlids that construct elaborate male display platforms, or bowers. Sister taxa with distinct bower morphologies, and that exist in discrete leks separated by only 1-2 m of depth, are divergent in both sexually selected and ecological traits. Our phylogeny suggests that the forces of sexual and ecological selection are intertwined during the speciation of this group and that specific bower characteristics and trophic morphologies have evolved repeatedly. These results suggest that trophic morphology and bower form may be inappropriate characters for delineating taxonomic lineages. Specifically the morphological characters used to describe the genera Lethrinops and Tramitichromis do not define monophyletic clades. Using a combination of behavioural and genetic characters, we were able to identify several cryptic cichlid species on a single beach, which suggests that sand dweller species richness has been severely underestimated.

Animals↗