Molecular and morphological characterization of Pyricularia and allied genera.

The phylogenetic relationships of Pyricularia species and species from related genera were established from sequences of the internal transcribed spacer ribosomal RNA gene. Phylogenetic analysis disclosed a consistent correlation with spore morphology. Most Pyricularia species studied, and two species of Dactylaria that have obpyriform conidia, fell within the Magnaporthaceae cluster with high bootstrap support. Pyricularia variabilis was more related to Dactylaria, Tumularia or Ochroconis species than to the Magnaporthaceae. Dactylaria and species of Nakataea, Ochroconis, Pyriculariopsis and Tumularia were distinct from the Magnaporthaceae, and the genus Dactylaria is polyphyletic. The combination of morphological and molecular characters, such as spore morphology and ITS ribosomal DNA sequences data, suggested that conidial shape could be a primary character to distinguish Pyricularia from related genera.

Bidirectional interlocus concerted evolution following allopolyploid speciation in cotton (Gossypium).

Polyploidy is a prominent process in plant evolution; yet few data address the question of whether homeologous sequences evolve independently subsequent to polyploidization. We report on ribosomal DNA (rDNA) evolution in five allopolyploid (AD genome) species of cotton (Gossypium) and species representing their diploid progenitors (A genome, D genome). Sequence data from the internal transcribed spacer regions (ITS1 and ITS2) and the 5.8S gene indicate that rDNA arrays are homogeneous, or nearly so, in all diploids and allopolyploids examined. Because these arrays occur at four chromosomal loci in allopolyploid cotton, two in each subgenome, repeats from different arrays must have become homogenized by interlocus concerted evolution. Southern hybridization analysis combined with copy-number estimation demonstrate that this process has gone to completion in the diploids and to completion or near-completion in all allopolyploid species and that it most likely involves the entire rDNA repeat. Phylogenetic analysis demonstrates that interlocus concerted evolution has been bidirectional in allopolyploid species--i.e., rDNA from four polyploid lineages has been homogenized to a D genome repeat type, whereas sequences from Gossypium mustelinum have concerted to an A genome repeat type. Although little is known regarding the functional significance of interlocus concerted evolution of homeologous sequences, this study demonstrates that the process occurs for tandemly repeated sequences in diploid and polyploid plants. That interlocus concerted evolution can occur bidirectionally subsequent to hybidization and polyploidization has significant implications for phylogeny reconstruction, especially when based on rDNA sequences.

An unusual ribosomal DNA sequence from Gossypium gossypioides reveals ancient, cryptic, intergenomic introgression.

The New World allopolyploid (AD-genome) cottons (Gossypium) originated through hybridization of ancestral diploid species that presently have allopatric ranges in Africa-Asia (the A-genome) and the American tropics and subtropics (the D-genome). Phylogenetic analysis of sequence data from the ribosomal DNA internal transcribed spacer region (ITS1, 5.8S rRNA, ITS2) reveals two strongly supported clades, one corresponding to African species and the other containing all American D-genome species except Gossypium gossypioides, which occupies a basal position within the African clade. This placement conflicts with evidence from fertility relationships, morphology, cytogenetics allozymes, and cpDNA restriction site analysis, which specify a sister-species relationship for G. gossypioides and Gossypium raimondii, deeply embedded within the American D-genome clade. Several alternative explanations for this striking incongruence are considered. The most probable involves: (1) an ancient hybridization event, whereby G. gossypioides experienced contact with an A-genome, either at the diploid level or at the triploid level as a consequence of hybridization with a New World allopolyploid and (2) repeated backcrossing of the hybrid into the G. gossypioides lineage, thereby restoring a nearly pure D-genome, but in the process generating a recombinant ribosomal DNA. We suggest that this process may implicate G. gossypioides, rather than G. raimondii, as the closest living descendant of the ancestral D-genome parent of the allopolyploids. In addition, this example of cryptic, intergenomic introgression between species groups now occupying different hemispheres illustrates how ITS data may provide insights into genome composition and evolutionary history, especially when used in the context of a holistic view that encompasses information from a variety of sources.

The tortoise and the hare: choosing between noncoding plastome and nuclear Adh sequences for phylogeny reconstruction in a recently diverged plant group.

Phylogenetic resolution is often low within groups of recently diverged taxa due to a paucity of phylogenetically informative characters. We tested the relative utility of seven noncoding cpDNA regions and a pair of homoeologous nuclear genes for resolving recent divergences, using tetraploid cottons (Gossypium) as a model system. The five tetraploid species of Gossypium are a monophyletic assemblage derived from an allopolyploidization event that probably occurred within the last 0.5-2 million years. Previous analysis of cpDNA restriction site data provided only partial resolution within this clade despite a large number of enzymes employed. We sequenced three cpDNA introns (rpl16, rpoC1, ndhA) and four cpDNA spacers (accD-psaI, trnL-trnF, trnT-trnL, atpB-rbcL) for a total of over 7 kb of sequence per taxon, yet obtained only four informative nucleotide substitutions (0.05%) resulting in incomplete phylogenetic resolution. In addition, we sequenced a 1.65-kb region of a homoeologous pair of nuclear-encoded alcohol dehydrogenase (Adh) genes. In contrast with the cpDNA sequence data, the Adh homoeologues yielded 25 informative characters (0.76%) and provided a robust and completely resolved topology that is concordant with previous cladistic and phenetic analyses. The enhanced resolution obtained using the nuclear genes reflects an approximately three- to sixfold increase in nucleotide substitution rate relative to the plastome spacers and introns.