title: Wolbachia transmission dynamics in Formica wood ants.
abstract: BACKGROUND
  The role of Wolbachia endosymbionts in shaping the mitochondrial diversity of their arthropod host depends on the effects they have on host reproduction and on the mode of transmission of the bacteria. We have compared the sequence diversity of wsp (Wolbachia surface protein gene) and the host mtDNA in a group of Formica ant species that have diverged approximately 0.5 million years ago (MYA). The aim was to study the relationship of Wolbachia and its ant hosts in terms of vertical and horizontal transmission of the bacteria.
  
  
  RESULTS
  All studied ant species were doubly infected with two Wolbachia strains (wFex1 and wFex4) all over their geographical distribution area in Eurasia. The most common haplotypes of these strains were identical with strains previously described from a more distantly related Formica ant, with an estimated divergence time of 3.5 - 4 MYA. Some strain haplotypes were associated to the same or closely related mtDNA haplotypes as expected under vertical transmission. However, in several cases the wsp haplotypes coexisted with distant mtDNA haplotypes, a pattern which is more compatible with horizontal transmission of the bacteria.
  
  
  CONCLUSIONS
  Two lines of evidence suggest that the sharing of Wolbachia strains by all F. rufa species is rather due to horizontal than vertical transmission. First, the fact that endosymbiont strains identical to those of F. rufa ants have been found in another species that diverged 3.5-4 MYA strongly suggests that horizontal transfer can and does occur between Formica ants. Second, the frequent sharing of identical Wolbachia strains by distant mitochondrial lineages within the F. rufa group further shows that horizontal transmission has occurred repeatedly. Nevertheless, our dataset also provides some evidence for longer-term persistence of infection, indicating that Wolbachia infection within this host clade has been shaped by both horizontal and vertical transmission of symbionts. The fact that all the ants were infected irrespective of the family structure of their societies gives no support to the proposed hypotheses that the spreading of Wolbachia in ants might be associated to the types of their societies.
authors.affiliation: Department of Biology, PO Box 3000, 90014 University of Oulu, Finland. lumi.viljakainen@oulu.fi
authors.forename: Lumi
authors.initials: L
authors.lastname: Viljakainen
authors.forename: Max
authors.initials: M
authors.lastname: Reuter
authors.forename: Pekka
authors.initials: P
authors.lastname: Pamilo
mesh_terms: D000818:Animals; D001000:Ants / Q000145:classification / Q000382:microbiology*; D001425:Bacterial Outer Membrane Proteins / Q000235:genetics; D004269:DNA, Bacterial / Q000235:genetics; D004272:DNA, Mitochondrial / Q000235:genetics; D019143:Evolution, Molecular; D022761:Gene Transfer, Horizontal; D014644:Genetic Variation; D006239:Haplotypes; D054884:Host-Pathogen Interactions; D017422:Sequence Analysis, DNA; D013045:Species Specificity; D013559:Symbiosis; D020577:Wolbachia / Q000145:classification / Q000502:physiology*
chemical_list: D001425:Bacterial Outer Membrane Proteins; D004269:DNA, Bacterial; D004272:DNA, Mitochondrial; C112171:Wsp protein, Wolbachia