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However, during walking, hypaxial muscles on both sides of the trunk are active at different points with the obliquus externus superficialis and profundus muscles active on the bending side, and the obliquus internus and transversalis muscles active on the side opposite the bending ( Carrier 1993 Bennett et al. During swimming, activation of epaxial muscles on one side of the trunk allows the body to create a concave curve contributing to the generation of a traveling wave ( Frolich and Biewener 1992 Carrier 1993). During swimming, salamanders propel themselves through undulations of the trunk and tail, passing a traveling wave down the body axis in contrast, walking salamanders rely heavily on their limbs for propulsion and generate a standing wave with their body axis ( Frolich and Biewener 1992).Įpaxial muscles are active to produce these motions during both swimming and walking but display distinct activation patterns in each behavior. Salamanders are commonly viewed as a model for studies of the terrestrial locomotion of early tetrapods because their sprawling limb posture and generalized body plan are thought to resemble those of many of the earliest vertebrates to move over land ( Edwards 1989 Ashley-Ross and Bechtel 2004). For example, one significant functional change that resulted from tetrapods becoming more terrestrial is the transfer of propulsive forces through the limbs and limb girdles to the vertebral column ( Kardong 2012). These anatomical changes to the axial skeleton reflect the different mechanical demands experienced by parts of the vertebral column as well as by different taxa ( Kardong 2012). As tetrapods became more terrestrial, the axial skeleton became more robust and different regions along the length of the vertebral column became specialized for different functional roles ( Long and Gordon 2004). Once early tetrapods began making forays onto land, significant changes to the musculoskeletal system were necessary to support their body weight out of the water and their enhanced terrestrial locomotion. These results suggest that tail undulations may be a more critical component to sprawling-postured tetrapod locomotion than previously recognized. Thus, salamanders respond to decreased lateral movement in their trunk by increasing movements in their tail, without changes in limb kinematics. There were no significant differences for any limb kinematic variables among treatments including average, minimum, and maximum angles.
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PVC and Tygon individuals had significantly higher curvature in the tails compared with unrestricted individuals. 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. 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. 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. 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). This reduction was performed by artificially limiting trunk flexibility by attaching a 2-piece shell around the body between the pectoral and pelvic girdles.
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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. The goal of this study was to quantify the effect of reduced lateral flexibility in a generalized sprawling tetrapod, the tiger salamander ( Ambystoma tigrinum). Despite their bony carapace preventing lateral undulations, turtles are able to improve their locomotor performance by increasing stride length via greater limb protraction. These benefits make them key characteristics of the locomotion of tetrapods with sprawling posture, with the exception of turtles. Lateral undulation and trunk flexibility offer performance benefits to maneuverability, stability, and stride length (via speed and distance traveled).