Although the red-legged running frog, Kassina maculata, is secondarily a walker/runner, it retains the capacity for multiple locomotor modes, including jumping at a wide range of angles (nearly 70 deg). Using simultaneous hind limb kinematics and singlefoot ground reaction forces, we performed inverse dynamics analyses to calculate moment arms and torques about the hind limb joints during jumping at different angles in K. maculata. We show that forward thrust is generated primarily at the hip and ankle, while body elevation is primarily driven by the ankle. Steeper jumps are achieved by increased thrust at the hip and ankle and greater downward rotation of the distal limb segments. Because of its proximity to the GRF vector, knee posture appears to be important in controlling torque directions about this joint and, potentially, torque magnitudes at more distal joints. Other factors correlated with higher jump angles include increased body angle in the preparatory phase, faster joint openings and increased joint excursion, higher ventrally directed force, and greater acceleration and velocity. Finally, we demonstrate that jumping performance in K. maculata does not appear to be compromised by presumed adaptation to walking/ running. Our results provide new insights into how frogs engage in a wide range of locomotor behaviours and the multi-functionality of anuran limbs.

The kinematic flexibility of frog hindlimbs enables multiple locomotor modes within a single species. Prior work has extensively explored maximum performance capacity in frogs; however, the mechanisms by which anurans modulate performance within locomotor modes remain unclear. We explored how Kassina maculata, a species known for both running and jumping abilities, modulates take-off angle from horizontal to nearly vertical. Specifically, how do 3D motions of leg segments coordinate to move the centre of mass (COM) upwards and forwards? How do joint rotations modulate jump angle? High-speed video was used to quantify 3D joint angles and their respective rotation axis vectors. Inverse kinematics was used to determine how hip, knee and ankle rotations contribute to components of COM motion. Independent of take-off angle, leg segment retraction (rearward rotation) was twofold greater than adduction (downward rotation). Additionally, the joint rotation axis vectors reoriented through time, suggesting dynamic shifts in relative roles of joints. We found two hypothetical mechanisms for increasing take-off angle. Firstly, greater knee and ankle excursion increased shank adduction, elevating the COM. Secondly, during the steepest jumps, the body rotated rapidly backwards to redirect the COM velocity. This rotation was not caused by pelvic angle extension, but rather by kinematic transmission from leg segments via reorientation of the joint rotation axes. We propose that K. maculata uses proximal leg retraction as the principal kinematic drive while dynamically tuning jump trajectory by knee and ankle joint modulation.

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