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Background   Galileo  is a transposable element responsible for the generation of three chromosomal inversions in natural populations of  Drosophila buzzatii . Although the most characteristic feature of  Galileo  is the long internally-repetitive terminal inverted repeats (TIRs), which resemble the  Drosophila Foldback  element, its transposase-coding sequence has led to its classification as a member of the  P-element  superfamily (Class II, subclass 1, TIR order). Furthermore,  Galileo  has a wide distribution in the genus  Drosophila , since it has been found in 6 of the 12  Drosophila  sequenced genomes. Among these species,  D. mojavensis , the one closest to  D. buzzatii , presented the highest diversity in sequence and structure of  Galileo  elements.    Results  In the present work, we carried out a thorough search and annotation of all the  Galileo  copies present in the  D. mojavensis  sequenced genome. In our set of 170  Galileo  copies we have detected 5  Galileo  subfamilies (C, D, E, F, and X) with different structures ranging from nearly complete, to only 2 TIR or solo TIR copies. Finally, we have explored the structural and length variation of the  Galileo  copies that point out the relatively frequent rearrangements within and between  Galileo  elements. Different mechanisms responsible for these rearrangements are discussed.    Conclusions  Although  Galileo  is a transposable element with an ancient history in the  D. mojavensis  genome, our data indicate a recent transpositional activity. Furthermore, the dynamism in sequence and structure, mainly affecting the TIRs, suggests an active exchange of sequences among the copies. This exchange could lead to new subfamilies of the transposon, which could be crucial for the long-term survival of the element in the genome.

Striking structural dynamism and nucleotide sequence variation of the transposon Galileo in the genome of Drosophila mojavensis