Turtling the Salamander: Tail Movements Mitigate Need for Kinematic Limb Changes during Walking in Tiger Salamanders (Ambystoma tigrinum) with Restricted Lateral Movement.

Abstract

Lateral undulation and trunk flexibility offer performance benefits to maneuverability, stability, and stride length (via speed and distance traveled). These benefits make them key characteristics of the locomotion of tetrapods with sprawling posture, with the exception of turtles. Despite their bony carapace preventing lateral undulations, turtles are able to improve their locomotor performance by increasing stride length via greater limb protraction. The goal of this study was to quantify the effect of reduced lateral flexibility in a generalized sprawling tetrapod, the tiger salamander (Ambystoma tigrinum). We had two potential predictions: (1) either salamanders completely compensate by changing their limb kinematics, or (2) their performance (i.e., speed) will suffer due to the reduced lateral flexibility. This reduction was performed by artificially limiting trunk flexibility by attaching a 2-piece shell around the body between the pectoral and pelvic girdles. Adult tiger salamanders (n = 3; SVL = 9-14.5 cm) walked on a 1-m trackway under three different conditions: unrestricted, flexible shell (Tygon tubing), and rigid shell (PVC tubing). Trials were filmed in a single, dorsal view, and kinematics of entire midline and specific body regions (head, trunk, tail), as well as the fore and hind limbs, were calculated. Tygon individuals had significantly higher curvature than both PVC and unrestricted individuals for the body, but this trend was primarily driven by changes in tail movements. PVC individuals had significantly lower curvature in the trunk region compared with unrestricted individuals or Tygon; however, there was no difference between unrestricted and Tygon individuals suggesting the shells performed as expected. PVC and Tygon individuals had significantly higher curvature in the tails compared with unrestricted individuals. There were no significant differences for any limb kinematic variables among treatments including average, minimum, and maximum angles. Thus, salamanders respond to decreased lateral movement in their trunk by increasing movements in their tail, without changes in limb kinematics. These results suggest that tail undulations may be a more critical component to sprawling-postured tetrapod locomotion than previously recognized.