ParagraphCentromeres are critical for cell division, loading CENH3/CENPA histone variant nucleosomes, directing kinetochore formation and allowing chromosome segregation 1,2 . Despite their conserved function, centromere size and structure are diverse across species. To understand this centromere paradox 3,4 , it is necessary to know how centromeric diversity is generated, and whether it reflects ancient trans-species variation or, instead, rapid post-speciation divergence. To address these questions, we assembled 346 centromeres from 66 Arabidopsis thaliana and two A. lyrata accessions, which revealed a remarkable degree of intra-and inter-species diversity. Arabidopsis thaliana centromere repeat arrays are embedded in linkage blocks, despite ongoing internal satellite turnover, demonstrating a role for unidirectional gene conversion in sequence diversification, rather than interhomolog unequal crossover. Additionally, centrophilic ATHILA transposons have recently invaded the satellite arrays. To counter ATHILA invasion, chromosome-specific bursts of satellite homogenization generate higher-order repeats and purge transposons, consistent with cycles of repeat evolution. Centromeric sequence changes are even more extreme in comparison between A. thaliana and A. lyrata. Together, our findings reveal rapid cycles of transposon invasion and purging via satellite homogenization, which drive centromere evolution and ultimately contribute to speciation.'centrophilic' (Fig. 3a-3b, Extended Data Fig. 7a-7b and Supplementary Table 5). The ATHILA 8