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Coevolution between a Family of Parasite Virulence Effectors and a Class of LINE-1 Retrotransposons ´ 1. 2. 3 1¤ Soledad Sacristan *, Marielle Vigouroux , Carsten Pedersen , Pari Skamnioti , Hans 3 4 1 1 Thordal-Christensen , Cristina Micali , James K. M. Brown , Christopher J. Ridout 1 Department of Disease and Stress Biology, John Innes Centre, Norwich, United Kingdom, 2 Computational and Systems Biology Department, John Innes Centre, Norwich, United Kingdom, 3 Department of Agriculture and Ecology, Faculty of Life Sciences, University of Copenhagen, Copenhagen, Denmark, 4 Department of Plant- ¨ Microbe Interactions, Max-Planck Institute for Plant Breeding Research, Koln, Germany Abstract Parasites are able to evolve rapidly and overcome host defense mechanisms, but the molecular basis of this adaptation is poorly understood. Powdery mildew fungi (Erysiphales, Ascomycota) are obligate biotrophic parasites infecting nearly 10,000 plant genera. They obtain their nutrients from host plants through specialized feeding structures known as haustoria. We previously identified the AVR k1 powdery mildew-specific gene family encoding effectors that contribute to the successful establishment of haustoria. Here, we report the extensive proliferation of the AVRk1 gene family throughout the genome of B. graminis, with sequences diverging in formae speciales adapted to infect different hosts. Also, importantly, we have discovered that the effectors have c