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How TE families evolve following expansion, specifically why distinct descendant lineages retain or lose mobilization competence, remains poorly understood. The PIF\/Harbinger superfamily, whose canonical elements encode two distinct proteins required for mobilization\u2014a DDE transposase and a MADF DNA-binding protein\u2014provides a compelling model to address this question. In Caenorhabditis inopinata, the closest known relative of C. elegans, we identified a spontaneous dumpy mutant caused by insertion of a PIF\/Harbinger element into the Cin-dpy-11 coding region. The inserted element, designated Harbinger-1M_cIno and belonging to the C. elegans Turmoil2 lineage, retains a MADF domain but lacks a recognizable DDE transposase open reading frame. Genome-wide curation recovered 258 related copies, revealing a strongly asymmetric family structure: noncoding and MADF-bearing derivatives expanded, whereas only a single locus retained an intact DDE gene. This demonstrates a novel functional partitioning where catalytic (DDE) and DNA-binding (MADF) roles are physically separated across distinct elements and supplied in trans. A second family, Harbinger-2M_cIno (1,376 copies), lacks a DDE source entirely and exhibits greater sequence divergence and terminal degradation. Together, these findings reveal that PIF\/Harbinger lineages in C. inopinata differ in their timing of expansion and mobilization competence, illustrating how bipartite TEs maintain activity through functional division while limiting host toxicity.<\/jats:p>","DOI":"10.1093\/gbe\/evag240","type":"journal-article","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T19:20:44Z","timestamp":1790623244000},"source":"Crossref","is-referenced-by-count":0,"title":["Active and Unusually Expanded\n                    <i>PIF\/Harbinger<\/i>\n                    Transposable Elements in the\n                    <i>Caenorhabditis inopinata<\/i>\n                    Genome"],"prefix":"10.1093","volume":"18","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-2182-6883","authenticated-orcid":false,"given":"Kazuki","family":"Sato","sequence":"first","affiliation":[{"name":"Laboratory of Parasite Systems Biology, Department of Integrated Biosciences, Graduate School of Frontier Sciences, The 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