Phylogeny and polyploidy: resolving the classification of cyprinine fishes (Teleostei: Cypriniformes).

Cyprininae is the largest subfamily (>1300 species) of the family Cyprinidae and contains more polyploid species (∼400) than any other group of fishes. We examined the phylogenetic relationships of the Cyprininae based on extensive taxon, geographical, and genomic sampling of the taxa, using both mitochondrial and nuclear genes to address the phylogenetic challenges posed by polyploidy. Four datasets were analyzed in this study: two mitochondrial gene datasets (465 and 791 taxa, 5604bp), a mitogenome dataset (85 taxa, 14,771bp), and a cloned nuclear RAG1 dataset (97 taxa, 1497bp). Based on resulting trees, the subfamily Cyprininae was subdivided into 11 tribes: Probarbini (new; Probarbus+Catlocarpio), Labeonini Bleeker, 1859 (Labeo & allies), Torini Karaman, 1971 (Tor, Labeobarbus & allies), Smiliogastrini Bleeker, 1863 (Puntius, Enteromius & allies), Poropuntiini (Poropuntius & allies), Cyprinini Rafinesque, 1815 (Cyprinus & allies), Acrossocheilini (new; Acrossocheilus & allies), Spinibarbini (new; Spinibarbus), Schizothoracini McClelland, 1842 (Schizothorax & allies), Schizopygopsini Mirza, 1991 (Schizopygopsis & allies), and Barbini Bleeker, 1859 (Barbus & allies). Phylogenetic relationships within each tribe were discussed. Two or three distinct RAG1 lineages were identified for each of the following tribes Torini, Cyprinini, Spinibarbini, and Barbini, indicating their hybrid origin. The hexaploid African Labeobarbus & allies and Western Asian Capoeta are likely derived from two independent hybridization events between their respective maternal tetraploid ancestors and Cyprinion.

Evolutionary history of Otophysi (Teleostei), a major clade of the modern freshwater fishes: Pangaean origin and Mesozoic radiation.

BACKGROUND: Freshwater harbors approximately 12,000 fish species accounting for 43% of the diversity of all modern fish. A single ancestral lineage evolved into about two-thirds of this enormous biodiversity (≈ 7900 spp.) and is currently distributed throughout the world's continents except Antarctica. Despite such remarkable species diversity and ubiquity, the evolutionary history of this major freshwater fish clade, Otophysi, remains largely unexplored. To gain insight into the history of otophysan diversification, we constructed a timetree based on whole mitogenome sequences across 110 species representing 55 of the 64 families. RESULTS: Partitioned maximum likelihood analysis based on unambiguously aligned sequences (9923 bp) confidently recovered the monophyly of Otophysi and the two constituent subgroups (Cypriniformes and Characiphysi). The latter clade comprised three orders (Gymnotiformes, Characiformes, Siluriformes), and Gymnotiformes was sister to the latter two groups. One of the two suborders in Characiformes (Characoidei) was more closely related to Siluriformes than to its own suborder (Citharinoidei), rendering the characiforms paraphyletic. Although this novel relationship did not receive strong statistical support, it was supported by analyzing independent nuclear markers. A relaxed molecular clock Bayesian analysis of the divergence times and reconstruction of ancestral habitats on the timetree suggest a Pangaean origin and Mesozoic radiation of otophysans. CONCLUSIONS: The present timetree demonstrates that survival of the ancestral lineages through the two consecutive mass extinctions on Pangaea, and subsequent radiations during the Jurassic through early Cretaceous shaped the modern familial diversity of otophysans. This evolutionary scenario is consistent with recent arguments based on biogeographic inferences and molecular divergence time estimates. No fossil otophysan, however, has been recorded before the Albian, the early Cretaceous 100-112 Ma, creating an over 100 million year time span without fossil evidence. This formidable ghost range partially reflects a genuine difference between the estimated ages of stem group origin (molecular divergence time) and crown group morphological diversification (fossil divergence time); the ghost range, however, would be filled with discoveries of older fossils that can be used as more reasonable time constraints as well as with developments of more realistic models that capture the rates of molecular sequences accurately.

Extensive hybridization and tetrapolyploidy in spined loach fish.

Hybridization and polyploidy are particular characteristics of biodiversity among cypriniform fish. To infer ancient parentage of the tetraploids found among Japanese Cobitis loaches, we compare mitochondrial and four nuclear loci from two tetraploids with several diploid populations of C. biwae, C. striata and related species. Mitochondrial gene tree showed monophyletic diploid C. striata and paraphyletic C. biwae relative to C. striata. On the contrary, nuclear sequence analysis recovered both monophyletic diploid C. biwae and C. striata. Disagreement in mitochondrial and nuclear gene trees indicates an ancient hybridization event and mitochondrial gene introgression. The tetraploid C. biwae includes both mitochondrial and nuclear sequences close to diploid C. biwae, indicating autotetraploidy. The tetraploid C. striata has nuclear alleles of both C. biwae and C. striata and mtDNA of C. biwae. It is an allotetraploid with C. biwae as the mother. Our analyses also reveal an onset of genome reshaping after the allotetraploidization.

Mitotic and meiotic analyses of the 'large race' of Cobitis striata, a polyploid spined loach of hybrid origin.

The large race of the Cobitis striata complex is a bisexual polyploid population. It is difficult to distinguish this polyploid loach morphologically from a sympatric diploid C. striata, the Biwa small race, indicating the close relationship between these two populations. The polyploid loach does have striata-specific satellite DNAs, but it also harbors C. biwae-related mtDNA. The large race has 2n=4x=98 chromosomes with 56 acrocentric chromosomes. Diploid C. striata has 2n=50 and C. Biwae 2n=48 chromosomes, suggesting the karyotype of polyploid striata as a combination of these two genomes. Meiotic figures showed a few quadrivalent formations besides many bivalents. Many of the quadrivalents were of metacentric chromosomes. Some were, however, made of four acrocentric or of two acro- plus two metacentric chromosomes. Chiasmata were visible in some quadrivalent association. Quadrivalent formation with chiasmata indicates the presence of homologous segments capable of crossing-over between two genomes. Thus, the general polyploid model is also applicable to this case of polyploidy.