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This approach had been pioneered in the classical work of Dobzhansky and Sturtevant who used inversions in Drosophila chromosomes to construct an evolutionary tree [ 12 ] . Subsequently, mathematical methods have been developed to calculate rearrangement distances between genomes, and, using these, phylogenetic trees have been built for certain small genomes, such as plant mitochondria and herpesviruses [ 13 14 ] . These approaches, however, are applicable only to genomes that show significant conservation of global gene order, which is manifestly not the case among prokaryotes [ 15 16 17 ] . Even relatively close species such as, for example, Escherichia coli and Haemophilus influenzae, two species of the γ-subdivision of Proteobacteria, retain very little conservation of gene order beyond the operon level (typically, two-to-four genes in a row), and essentially none is detectable among evolutionarily distant bacteria and archaea [ 15 16 18 ] . Very few operons, primarily those coding for physically interacting subunits of multiprotein complexes such as certain ribosomal proteins or RNA-polymerase subunits, are conserved across a wide range of prokaryotic lineages [ 15 16 ] . On the other hand, pairwise comparisons of even distantly related prokaryotic genomes reveal considerable number of shared (predicted) operons, which creates an opportunity for a meaningful comparative analysis [ 19 ] [ 20 21 ] .
Reinforcement, a concept popularized by Dobzhansky (1937), is a process by which speciation, a macroevolutionary process, can be driven directly by natural selection, one of the primary microevolutionary forces.
, lack of mating or zygote formation) between species and thereby reduce production of unfit hybrids, even if Mayr did not share Dobzhansky's belief that this was the norm.
This process, the evolution of premating isolation after secondary contact due to selection against hybrids, is reinforcement sensu Dobzhansky (1937).
From the seminal population-genetic and phylogenetic studies of Dobzhansky and co-workers [ 1], and the classification of taxonomic relationships in the genus by Patterson, Stone and others [ 2], Drosophila has long served as a model system for developing and testing evolutionary principles at the morphological and cytological levels.
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