Kinematic analysis of tongue movements during chemosensory behaviour in the European green lizard, Lacerta viridis (Reptilia: Lacertidae)

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Abstract
High-speed cinematography (100 frames/s) was used to allow quantitative analysis of the kinematic profiles of tongue and jaw displacements during chemosensory activities in the scleroglossan lizard Lacerta viridis. The types of tongue flicking were simple downward extensions (SDE), single oscillations (SOC), and submultiple oscillations (SMOC) of the tongue out of the mouth. The SMOC type involves a downward or upward movement of the tongue performed before a typical oscillation and it is therefore suggested that this is an intermediate category of flick between the typical SOC and MOC of lizards. Closing and opening of the mouth in SDE, SOC, and SMOC cycles may or may not be separated by a stationary stage during which the jaws are held open at a constant gape. The duration of this stationary interval increases from SDE to SMOC. Gape cycles do not show any division into slow and fast stages. The gape is produced largely by depression of the lower jaw; the upper jaw is slightly elevated by protrusion of the tongue. Patterns of correlation of kinematic variables depicting jaw and tongue movements differed between SDE, SOC, and SMOC. A principal component analysis shows that the three flick types overlap in a multivariate space constructed from the kinematic variables depicting jaw and tongue displacements. Overlap between SOC and SMOC categories is greater than that between SOC, SMOC, and SDE categories. The kinematic patterns of tongue displacement during SMOC in Lacerta viridis show similarities with those of MOC in other lizards and in snakes. Kinematically, the pattern of jaw and tongue displacements of Lacerta viridis during chemosensory activities shows similarities with those that occur during drinking and prey capture.

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