“… Graham, 1973 ; Fretter and Graham, 1976 ; Crampton, 1977 ; Morse, 1984 ; Simone, 2005 , 2011 ; Evans et al, 2009 ; Golding et al, 2009 ; Balog et al, 2012 ) and the analysis of feeding tracks (e.g. Ankel, 1936 , 1938 ; Eigenbrodt, 1941 ; Hickman and Morris, 1985 ; Janssen and Triebskorn, 1987 ; Mackenstedt and Märkel, 2001 ) demonstrate that the motion apparatus is highly diverse and complex. Traditionally the radula was compared to a flat ribbon or a ‘belt-and-pulley’ model ( Ankel, 1938 ; Märkel, 1964 ; Solem, 1974 ; Hickman, 1980 ), but this view has subsequently changed.…”