Husnik F, McCutcheon JPPlease use the format "First name initials family name" as in "Marie S. Curie, Niels H. D. Bohr, Albert Einstein, John R. R. Tolkien, Donna T. Strickland"
Stable endosymbiosis of a bacterium into a host cell promotes cellular and genomic complexity. The mealybug *Planococcus citri* has two bacterial endosymbionts with an unusual nested arrangement: the γ-proteobacterium *Moranella endobia* lives in the cytoplasm of the β-proteobacterium *Tremblaya princeps*. These two bacteria, along with genes horizontally transferred from other bacteria to the *P. citri* genome, encode gene sets that form an interdependent metabolic patchwork. Here, we test the stability of this three-way symbiosis by sequencing host and symbiont genomes for five diverse mealybug species and find marked fluidity over evolutionary time. Although *Tremblaya* is the result of a single infection in the ancestor of mealybugs, the γ-proteobacterial symbionts result from multiple replacements of inferred different ages from related but distinct bacterial lineages. Our data show that symbiont replacement can happen even in the most intricate symbiotic arrangements and that preexisting horizontally transferred genes can remain stable on genomes in the face of extensive symbiont turnover.
scale insect, organelle, symbiosis, Horizontal Gene Transfer
