© 2019, American Society for Microbiology. The attached document (embargoed until 18/10/2019) is an author produced version of a paper published in APPLIED AND ENVIRONMENTAL MICROBIOLOGY uploaded in accordance with the publisher’s self- archiving policy. The final published version (version of record) is available online at the link. Some minor differences between this version and the final published version may remain. We suggest you refer to the final published version should you wish to cite from it.