Animal Mitochondria, Positive Selection and Cyto-Nuclear Coevolution Insights from Pulmonates.docVIP

AnimalMitochondria,PositiveSelectionandCyto- NuclearCoevolution:InsightsfromPulmonates AristeidisParmakelis1*,PanayiotaKotsakiozi2,DavidRand3 1Department of Ecology and Taxonomy, Faculty of Biology, National and Kapodistrian University of Athens, Athens, Greece, 2Department of Animal and Human Physiology, Faculty of Biology, National and Kapodistrian University of Athens, Athens, Greece, 3Department of Ecology and Evolutionary Biology, Brown University, Providence,RhodeIsland,UnitedStatesofAmerica Abstract Pulmonate snails have remarkably high levels of mtDNA polymorphism within species and divergence between species, makingthemaninterestinggroupforthestudyofmutationandselectiononmitochondrialgenomes.Theavailabilityof sequencedatafrommostmajor lineages–collectedlargelyforstudiesofphylogeography-provides anopportunity to performseveraltestsofselectionthatmayprovidegeneralinsightsintotheevolutionaryforcesthathaveproducedthis unusualpattern.SeveralproteincodingmtDNAdatasetsofpulmonateswereanalyzedtowardsthisdirection.Twodifferent methods for the detection of positive selection were used, one based on phylogeny, and the other on the McDonald- Kreitman test. The cyto-nuclear coevolution hypothesis, often implicated to account for the high levels of mtDNA divergenceofsomeorganisms,wasalsoaddressedbyassessingthedivergencepatternexhibitedbyanucleargene.The McDonald-Kreitman test indicated multiple signs of positive selection in the mtDNA genes, but was significantly biased when sequence divergence was high. The phylogenetic method identified five mtDNA datasets as affected by positive selection. In the nuclear gene, the McDonald-Kreitman test provided no significant results, whereas the phylogenetic methodidentifiedpositiveselectionaslikelypresent.Overall,ourfindingsindicatethat:1)slimsupportforthecyto-nuclear coevolution hypothesis is present, 2) the elevated rates of mtDNA polymorphims and divergence in pulmonates do not appeartobeduetopervasivepositiveselection,