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OATAO is an open access repository that collects the work of Toulouse researchers and makes it freely available over the web where possible. This is an author-deposited version published in : http://oatao.univ-toulouse.fr/ Eprints ID : 10815 To link to this article :doi:10.1099/ijs.0.039099-0 pmid:22523169 fatcat:pycecmplifartgzcnr53nxmrti