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Background   Tc1/mariner  transposons are widespread DNA transposable elements (TEs) that have made important contributions to the evolution of host genomic complexity in metazoans. However, the evolution and diversity of the  Tc1/mariner  superfamily remains poorly understood. Following recent developments in genome sequencing and the availability of a wealth of new genomes,  Tc1/mariner  TEs have been identified in many new taxa across the eukaryotic tree of life. To date, the majority of studies focussing on  Tc1 / mariner  elements have considered only a single host lineage or just a small number of host lineages. Thus, much remains to be learnt about the evolution of  Tc1/mariner  TEs by performing analyses that consider elements that originate from across host diversity.    Results  We mined the non-redundant database of NCBI using BLASTp searches, with transposase sequences from a diverse set of reference  Tc1/mariner  elements as queries. A total of 5158  Tc1/mariner  elements were retrieved and used to reconstruct evolutionary relationships within the superfamily. The resulting phylogeny is well resolved and includes several new groups of  Tc1/mariner  elements. In particular, we identify a new family of plant-genome restricted  Tc1/mariner  elements, which we call  PlantMar . We also show that the  pogo  family is much larger and more diverse than previously appreciated, and we review evidence for a potential revision of its status to become a separate superfamily.    Conclusions  Our study provides an overview of  Tc1-mariner  phylogeny and summarises the impressive diversity of  Tc1-mariner  TEs among sequenced eukaryotes.  Tc1/mariner  TEs are successful in a wide range of eukaryotes, especially unikonts (the taxonomic supergroup containing Amoebozoa, Opisthokonta, Breviatea, and Apusomonadida). In particular, ecdysozoa, and especially arthropods, emerge as important hosts for  Tc1/mariner  elements (except the  PlantMar  family). Meanwhile, the  pogo  family, which is by far the largest  Tc1/mariner  family, also includes many elements from fungal and chordate genomes. Moreover, there is evidence of the repeated exaptation of  pogo  elements in vertebrates, including humans, in addition to the well-known example of  CENP-B . Collectively, our findings provide a considerable advancement in understanding of  Tc1/mariner  elements, and more generally they suggest that much work remains to improve understanding of the diversity and evolution of DNA TEs.

Phylogenetic analysis of the Tc1/mariner superfamily reveals the unexplored diversity of pogo-like elements