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PubMed2006

Life history evolution in cichlids 2: directional evolution of the trade-off between egg number and egg size.

The negative relationship between offspring number and offspring size provides a classic example of the role of trade-offs in life history theory. However, the evolutionary transitions in egg size and clutch size that have produced this negative relationship are still largely unknown. Since body size may affect both of these traits, it would be helpful to understand how evolutionary changes in body size may have facilitated or constrained shifts in clutch and egg size. By using comparative methods with a database of life histories and a phylogeny of 222 genera of cichlid fishes, we investigated the order of evolutionary transitions in these traits in relation to each other. We found that the ancestral large-bodied cichlids first increased egg size, followed by a decrease in both body size and clutch size resulting in the common current combination of a small-bodied cichlid with a small clutch of large eggs. Furthermore, lineages that deviated from the negative relationship between clutch and egg size underwent different transitions in these traits according to their body size (large bodied genera have moved towards the large clutch/small egg end of the continuum and small bodied genera towards the small clutch/large egg end of the continuum) to reach the negative relationship between clutch size and egg size. Our results show that body size is highly important in shaping the negative relationship between clutch size and egg size.

Animals↗

PubMed2006

Colour forms of Amazonian cichlid fish represent reproductively isolated species.

Laboratory mate choice experiments have confirmed species status for cichlid fish in the African Great Lakes that differ in colour and little else. Colour differences between allopatric populations of the South American cichlid genus Apistogramma are known for many species, yet the status of such populations has not been previously tested. Analysis of the genetic relationships and mate choice characteristics of populations previously described as Apistogramma caetei from eastern Amazonia indicates genetic differentiation into at least three allopatric lineages, which also show strong prezygotic isolation through female mate choice, confirming them as Biological species. If future studies confirm that this result is indicative of a general trend, the species richness of the South American cichlid fishes may presently be seriously underestimated.

Animals↗

PubMed2001

Mechanisms of rapid sympatric speciation by sex reversal and sexual selection in cichlid fish.

Mechanisms of speciation in cichlid fish were investigated by analyzing population genetic models of sexual selection on sex-determining genes associated with color polymorphisms. 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 models explain why sex-reversal genes that change males into females tend to be X-linked and associated with novel colors, using the hypothesis of restricted recombination on the sex chromosomes, as suggested by previous theory on the evolution of recombination. The models reveal multiple pathways for rapid sympatric speciation through the origin of novel color morphs with strong assortative mating that incorporate both sex-reversal and suppressor genes. Despite the lack of geographic isolation or ecological differentiation, the new species coexists with the ancestral species either temporarily or indefinitely. These results may help to explain different patterns and rates of speciation among groups of cichlids, in particular the explosive diversification of rock-dwelling haplochromine cichlids.

Animals↗

PubMed1987

Coexpression of distinct eye- and liver-specific LDH isozymes in cichlid fish.

In a recent electrophoretic survey of lactate dehydrogenase (LDH) in neotropical cichlid fishes (Perciformes, Cichlidae) we have discovered several species in which a cathodal liver-specific isozyme is expressed along with the highly-anodal eye-specific isozyme (LDH-C4) typically encountered in perciform fishes. We believe this fourth, liver-specific LDH isozyme to be real and not artifactual since homogenization of fresh liver from one of these species, the Basketmouth cichlid (Acaronia nassa), in either of two nondenaturing detergents or in the presence of the protease inhibitor phenylmethylsulfonylfluoride affects neither the presence nor mobility of this cathodal band. Moreover, it continues to be expressed in the captively bred F1 of these same wild fish. The discovery of several fish species, like the Basketmouth, in which biochemically distinct eye- and liver-specific LDH isozymes are coexpressed, is discussed in light of the currently accepted hypothesis that these two isozymes are encoded by a single locus (LDH-C) which has undergone divergent tissue expression in several other major teleost groups. Preliminary characterization of the liver-specific isozyme relative to the eye-specific LDH-C4 in the Basketmouth cichlid with respect to thermolability and NADH-induced binding to oxamate-sepharose columns suggests that the eye- and liver-specific LDH isozymes are biochemically quite distinct in this fish and that they are probably encoded by two distinct loci.

Animals↗

PubMed1994

The utility of SATA satellite DNA sequences for inferring phylogenetic relationships among the three major genera of tilapiine cichlid fishes.

The SATA satellite DNA family of sequences, composed of three size variants of approximately 237, 230, and 209 bp, is conserved in the genomes of tilapiine and haplochromine cichlid fishes. In the present study we examined the utility of the SATA sequences for inferring phylogenetic relationships among the three major genera of tilapiine fishes, Oreochromis, Sarotherodon, and Tilapia. Hybridization of the monomer SATA repeat to genomic DNA of representative cichlid species established conservation of the sequence in the African tilapine and haplochromine lineages and its absence from other cichlid lineages. Bootstrapped DNA parsimony and neighbor-joining analyses of derived consensus sequences revealed two distinct clades, one containing the mouthbrooding genera Oreochromis and Sarotherodon, and the other containing the substrate spawning genus Tilapia. These results are consistent with recent independent studies using mitochondrial DNA and establish the utility of the SATA satellite DNA family for phylogenetic reconstruction. Concerted evolution of the SATA sequences was also demonstrated within the tilapiine tribe.

Animals↗

PubMed1995

Evolution of NADH dehydrogenase subunit 2 in east African cichlid fish.

The complete sequence (1047 bp) of the mitochondrially encoded ND2 gene was obtained from 31 species of cichlid fishes to investigate the evolutionary history of the species flocks of the East African lakes. The observed pattern of nucleotide substitution in this sequence is typical of mitochondrial genes, showing a high transition bias and rapid mutational saturation, especially at codon positions where base frequencies are unequal. The base composition of the third position of codons is heterogeneous among species, suggesting frequent shifts in the pattern of substitution. Phylogenetic analysis of the sequences shows that the mtDNA variation in Lake Malawi cichlids is nested monophyletically within the range of variation shown by Tanganyikan cichlids. The closest Tanganyikan relatives of the Malawi flock are members of the tribe Tropheini. Classifications based on morphology are generally supported by the mtDNA data, with some significant exceptions in the Tropheini and Lamprologini. Because of an apparently rapid radiation of the Tanganyikan lineages, it is difficult to assess the basal topology of the Tanganyikan radiation at this time. Divergences among tribes are consistent with an intralacustrine radiation within the past 10 million years.

Africa, Eastern↗

PubMed2000

Linkage Relationships and Haplotype Variation of the Major Histocompatibility Complex Class I A Genes in the Cichlid Fish Oreochromis niloticus.

The haplochromine cichlid species flocks of the East African Great Lakes are one of the best examples of adaptive radiation. Analysis of genetic variation among these species provides valuable information on species relationships and timing of speciation events. Although the haplochromine cichlids generally display little genetic variation, the major histocompatibility complex (Mhc) genes have been found to be highly variable. A study of the linkage relationships of the Mhc class I A genes in the cichlid fish Oreochromis niloticus was therefore undertaken. Class I loci were identified, and their segregation in seven mothers and their haploid embryos was determined. In total, 56 class I A sequences were found among the seven families. A strong concordance of segregation was observed in five haplotypes among the embryos, indicating a close linkage of all loci. The number of loci per haplotype varied from 11 to 17, while the total number of distinct loci found among all families was 22. These findings show that all class I A loci are linked in a single genetic cluster in O. niloticus.

Journal Article↗

PubMed1997

Class I mhc genes of cichlid fishes: identification, expression, and polymorphism.

Cichlid fishes of the East African Rift Valley lakes constitute an important model of adaptive radiation. Explosive speciation in the Great Lakes, in some cases as recently as 12 400 years ago, generated large species flocks that have been the focus of evolutionary studies for some time. The studies have, however, been hampered by the paucity of biochemical markers for phylogenetic reconstruction. Here, we describe a set of markers which should help to alleviate this problem. They are the class I genes of the major histocompatibility complex. We provide evidence for the existence of at least 17 class I loci in cichlid fishes, and for extensive polymorphism of three of these loci. Since the polymorphism has a trans-species character, it will be possible to use it in investigating the founding events of the individual species. The sequences of the cichlid class I fishes support the monophyly of actinopterygian fish on the one hand, and of tetrapods on the other.

Animals↗

PubMed2001

New markers for new species: microsatellite loci and the East African cichlids.

The radiation of the East African cichlid fishes has engaged biologists for over a century. Because so much taxonomic diversity has evolved recently, they are an ideal natural system in which to study the process of speciation. Hypervariable microsatellite loci have been used to verify multiple paternity and maternity in cichlid broods, to quantify the fitness of cooperative breeders and reproductive parasites, to estimate effective population sizes in captive populations, and to illuminate the spatial and temporal scale of gene flow among natural populations. The patterns that have emerged from these studies often reflect important biological differences among taxa. The cichlid species of East Africa represent a large amount of taxonomic and adaptive diversity all neatly packaged into a single lineage and confined to a modest geographical area. Data from microsatellite loci are now providing us with the means to understand one of the world's most intriguing and instructive comparative evolutionary systems.

Journal Article↗

PubMed1992

Genetic divergence, speciation and morphological stasis in a lineage of African cichlid fishes.

Since their discovery at the turn of the century, the species assemblages of cichlid fishes in the East African Lakes Victoria, Malawi and Tanganyika have fascinated evolutionary biologists. Many models have attempted to account for the 'explosive' evolution of several hundred species within these lakes. Here we report a case of surprisingly large genetic divergence among populations of the endemic Tropheus lineage of Lake Tanganyika. This lineage of six species contains twice as much genetic variation as the entire morphologically highly diverse cichlid assemblage of Lake Malawi and six times more variation than the Lake Victoria species flock. Although it is highly variable in coloration, this group of species and its closest relatives have not undergone appreciable morphological change. The observed geographic pattern of genetic variation suggests that major lake level fluctuations affected the distribution and speciation of this lineage of cichlid fishes.

Africa↗

PubMed1994

Sympatric speciation suggested by monophyly of crater lake cichlids.

The existence of sympatric speciation--that populations diverge into species in the absence of physical or ecological barriers--is controversial. The East African Great Lakes harbour hundreds of cichlid species representing only a few monophyletic lineages, although palaeolimnological evidence and local restrictions on species distribution suggest that speciation in these lakes could have been allopatric. The case for sympatry in restricted areas of Lakes Malawi and Tanganyika is stronger but not unassailable. A better case might be made for cichlid species flocks in small, ecologically monotonous crater lakes. Here we present a mitochondrial DNA analysis of cichlid species flocks endemic to two such lakes in Cameroon. The results suggest that the flocks in each lake are monophyletic: the implication being that each lake was colonized once only, the size and shape of each lake being such that subsequent diversification would have been sympatric.

Animals↗

PubMed1975

Trophic radiation through polymorphism in cichlid fishes.

Several morphologically defined species of cichlid fishes (Cichlasoma) endemic to the Cuatro Cienegas basin of Mexico and differing in tooth structure, body shape, and diet are allelically identical at 27 gene loci. The presence of only one Mendelian population in each of three drainage systems studied and the occurrence of two of the morphotypes in the same broods indicate that the supposed species are morphs. That trophic radiation in the Cuatro Cienegas cichlids has been achieved through ecological polymorphism rather than speciation raises questions regarding the genetic basis for the extensive intralacustrine radiation of cichlids in Africa and elsewhere.

Journal Article↗

PubMed1999

Replicated evolution of trophic specializations in an endemic cichlid fish lineage from Lake Tanganyika.

The current phylogenetic hypothesis for the endemic Lake Tanganyika cichlid fishes of the tribe Eretmodini is based solely on morphology and suggests that more complex trophic morphologies derived only once from a less specialized ancestral condition. A molecular phylogeny of eretmodine cichlids based on partial mitochondrial DNA cytochrome b and control-region sequences was used to reconstruct the evolutionary sequence of trophic adaptations and to test alternative models of morphological divergence. The six mitochondrial lineages found disagree with the current taxonomy and the morphology-based phylogeny. Mitochondrial lineages with similar trophic morphologies are not grouped monophyletically but are typically more closely related to lineages with different trophic phenotypes currently assigned to other genera. Our results indicate multiple independent origins of similar trophic specializations in these cichlids. A pattern of repeated divergent morphological evolution becomes apparent when the phylogeography of the mitochondrial haplotypes is analyzed in the context of the geological and paleoclimatological history of Lake Tanganyika. In more than one instance within Lake Tanganyika, similar morphological divergence of dentitional traits occurred in sympatric species pairs. Possibly, resource-based divergent selective regimes led to resource partitioning and brought about similar trophic morphologies independently and repeatedly.

Animals↗

PubMed2000

Genetic population structure indicates sympatric speciation of Lake Malawi pelagic cichlids.

Allopatric processes of speciation have routinely been presented to explain the extraordinary radiation of the East African Great Lakes cichlid fish species flocks. The 21 or more species of pelagic cichlids within the Lake Malawi flock appear to have lake-wide distributions that challenge such a concept. Data from six microsatellite DNA loci indicate single, panmictic populations across the lake of three Diplotaxodon species. Levels of variability at these loci suggest that populations have been large and stable. Mitochondrial DNA sequence data (872 bp of control region + 981 bp of the NADH-2) from 90 species, representing all major clades within the Lake Malawi flock, indicate reciprocal monophyly of the pelagic clade. We suggest that these data support a hypothesis that speciation in sympatry is more plausible (and widespread) within the cichlid species flocks than previously thought.

Animals↗

PubMed1996

Mitochondrial phylogeography of rock-dwelling cichlid fishes reveals evolutionary influence of historical lake level fluctuations of Lake Tanganyika, Africa.

The East African Lakes Tanganyika, Malawi and Victoria each harbour hundreds of endemic invertebrate and vertebrate species. Inferences about the ecological and evolutionary processes responsible for the origin of these species flocks will only be possible when they are made within historical and comparative frameworks. Specifically, the relative importance of intrinsic characteristics and extrinsic factors may offer information about the processes that drive diversification and speciation in these species. We investigated the sequence variation of a segment of the mitochondrial DNA control region of 32 populations representing all four nominal species in the three genera of eretmodine cichlids from Lake Tanganyika. Based on a phylogenetic analysis of these data we attempted to evaluate the importance of major lake level fluctuations on patterns of intralacustrine speciation. The geography of genetic variation reveals a high degree of within-lake endemism among genetically well-separated lineages distributed along the inferred shore lines of three historically intermittent lake basins. Seismic data indicate that extreme lowering of water levels in the Pleistocene caused the single Lake Tanganyika basin to split into three isolated ones. The strong phylogeographic structure of the Eretmodini, and the observation that some closely related populations occur on opposite shores of the lake, agree with this geological scenario. The three-clade-three-basin phylogeographic pattern was repeated twice within this tribe of cichlids. The phylogeographic pattern of eretmodine cichlids suggests that major fluctuations in the level of the lake have been important in shaping their adaptive radiation and speciation. The mitochondrially defined clades are in conflict with the current taxonomy of the group and suggest that there has been convergent evolution in trophic morphology, particularly in the shapes of oral teeth, taxonomically the most diagnostic characters of the three genera.

Age Factors↗

PubMed1997

Microhabitat use, trophic patterns, and the evolution of brain structure in African cichlids.

The species assemblages of cichlids in the three largest African Great Lakes are among the richest concentrations of vertebrate species on earth. The faunas are broadly similar in terms of trophic diversity, species richness, rates of endemism, and taxonomic composition, yet they are historically independent of each other. Hence, they offer a true and unique evolutionary experiment to test hypotheses concerning the mutual dependencies of ecology and brain morphology. We examined the brains of 189 species of cichlids from the three large lakes: Victoria, Tanganyika, and Malawi. A first paper demonstrated that patterns of evolutionary change in cichlid brain morphology are similar across taxonomic boundaries as well as across the three lakes [van Staaden et al., 1995 ZACS 98: 165-178]. Here we report a close relationship between the relative sizes of various brain structures and variables related to the utilization of habitat and prey. Causality is difficult to assign in this context, nonetheless, prey size and agility, turbidity levels, depth, and substrate complexity are all highly predictive of variation in brain structure. Areas associated with primary sensory functions such as vision and taste relate significantly to differences in feeding habits. Turbidity and depth are closely associated with differences in eye size, and large eyes are associated with species that pick plankton from the water column. Piscivorous taxa and others that utilize motile prey are characterized by a well developed optic tectum and a large cerebellum compared to species that prey on molluscs or plants. Structures relating to taste are well developed in species feeding on benthos over muddy or sandy substrates. The data militated against the existence of compensatory changes in brain structure. Thus enhanced development of a particular function is generally not accompanied by a parallel reduction of structures related to other modalities. Although genetic and environmental influences during ontogeny of the brain cannot be isolated, this study provides a rich source of hypotheses concerning the way the nervous system functions under various environmental conditions and how it has responded to natural selection.

Africa↗

PubMed1998

ROn-1 SINEs: a tRNA-derived, short interspersed repetitive DNA element from Oreochromis niloticus and its species-specific distribution in Old World cichlid fishes.

A SINE-like repetitive element (ROn-1) has been cloned from the tilapiine cichlid fish Oreochromis niloticus. The element is 345 base pairs (bp) in length and consists of a transfer-RNA-like domain with putative RNA polymerase III recognition sequences, a tRNA-unrelated region, and a poly(A) tail. Approximately 6000 copies of ROn-1 occur in the haploid genome of O. niloticus. Southern blot analysis revealed that ROn-1 is an abundant element in the genomes of many African cichlid fishes, but absent from the genome of the Indian cichlid Etroplus.

Africa↗

PubMed2006

From conical to spatulate: intra- and interspecific changes in tooth shape in closely related cichlids (Teleostei; Cichlidae: Eretmodini).

The Eretmodini are closely related cichlids endemic to Lake Tanganyika with very divergent oral tooth shapes, ranging from spatulate in Eretmodus to conical in Tanganicodus. To study how closely related cichlids can generate such divergent tooth shapes, we investigated how the enamel organ directs the development of spatulate teeth in Eretmodus cf. cyanostictus (lineage A), both in ontogeny and in adults, and of conical teeth in adult Tanganicodus cf. irsacae, using 3D-reconstructions from serially sectioned tooth germs. The spatulate oral tooth shape that characterizes adult E. cf. cyanostictus (lineage A) is preceded early in ontogeny by a conical tooth shape. We propose two possible hypotheses to account for changes in the folding of the enamel organ (in particular its epithelio-mesenchymal boundary) capable of generating such distinct tooth shapes. Different arguments lead us to favor the hypothesis of an asymmetric growth and differentiation of the enamel organ, such that the tip of a conical tooth corresponds to one "corner" of a spatulate tooth. Applying current molecular models of tooth shape variation, this would imply the existence of asymmetric fields of inhibition. Whether such asymmetric growth reflects the reutilization of a simple mechanism operating in ontogeny has to be clarified.

Adaptation, Biological↗