Duplication and concerted evolution of the mitochondrial control region in the parrot genus Amazona

We report a duplication and rearrangement of the mitochondrial genome involving the control region of parrots in the genus Amazona. This rearrangement results in a gene order of cytochrome b/tRNA(Thr)/pND6/pGlu/CR1/tRNA(Pro)/NADH dehydrogenase 6/tRNA(Glu)/CR2/tRNA(Phe)/12s rRNA, where CR1 and CR2 refer to duplicate control regions, and pND6 and pGlu indicate presumed pseudogenes. In contrast to previous reports of duplications involving the control regions of birds, neither copy of the parrot control region shows any indications of degeneration. Rather, both copies contain many of the conserved sequence features typically found in avian control regions, including the goose hairpin, TASs, the F, C, and D boxes, conserved sequence box 1 (CSB1), and an apparent homolog to the mammalian CSB3. We conducted a phylogenetic analysis of homologous portions of the duplicate control regions from 21 individuals representing four species of Amazona (A. ochrocephala, A. autumnalis, A. farinosa, and A. amazonica) and Pionus chalcopterus. This analysis revealed that an individual's two control region copies (i.e., the paralogous copies) were typically more closely related to one another than to corresponding segments of other individuals (i.e., the orthologous copies). The average sequence divergence of the paralogous control region copies within an individual was 1.4%, versus a mean value of 4.1% between control region orthologs representing nearest phylogenetic neighbors. No differences were found between the paralogous copies in either the rate or the pattern in which the two copies accumulated base pair changes. This pattern suggests concerted evolution of the two control regions, perhaps through occasional gene conversion events. We estimated that gene conversion events occurred on average every 34,670 +/- 18,400 years based on pairwise distances between the paralogous control region sequences of each individual. Our results add to the growing body of work indicating that under some circumstances duplicated mitochondrial control regions are retained through evolutionary time rather than degenerating and being lost, presumably due to selection for a small mitochondrial genome.