The Gram reaction was determined using the bioMe´rieux Gram-stain kit, according to the manufacturer’s instructions. Gliding motility was investigated by phase-contrast light microscopy with the hanging drop method, as described by Bernardet et al. (2002). Anaerobic growth was tested on MA incubated for up to 15 days in the AnaeroPack-Anaero system (Mitsubishi Gas Chemical). Catalase and oxidase activities were assessed using standard methods (Tindall et al., 2007). The degradation of agar,DNA, starch and CM-cellulose was assessed according to the methods of Smibert & Krieg (1994).