Abstract

In bats, the wing membrane is anchored not only to the body and forelimb, but also to the hindlimb. This attachment configuration gives bats the potential to modulate wing shape by moving the hindlimb, such as by joint movement at the hip or knee. Such movements could modulate lift, drag, or the pitching moment. In this study we address: 1) how the ankle translates through space during the wingbeat cycle; 2) whether amplitude of ankle motion is dependent upon flight speed; 3) how tension in the wing membrane pulls the ankle; and 4) whether wing membrane tension is responsible for driving ankle motion. We flew five individuals of the lesser dog-faced fruit bat, Cynopterus brachyotis (Family: Pteropodidae), in a wind tunnel and documented kinematics of the forelimb, hip, ankle, and trailing edge of the wing membrane. Based on kinematic analysis of hindlimb and forelimb movements, we found that: 1) during downstroke, the ankle moved ventrally and during upstroke the ankle moved dorsally; 2) there was considerable variation in amplitude of ankle motion, but amplitude did not correlate significantly with flight speed; 3) during downstroke, tension generated by the wing membrane acted to pull the ankle dorsally, and during upstroke, the wing membrane pulled laterally when taut and dorsally when relatively slack; and 4) wing membrane tension generally opposed dorsoventral ankle motion. We conclude that during forward flight in C. brachyotis, wing membrane tension does not power hindlimb motion; instead, we propose that hindlimb movements arise from muscle activity and/or inertial effects.

Highlights

  • Bats are well known for their forelimbs, which are significantly modified into wings, but their hindlimbs are modified as well

  • We propose that hindlimb motion could arise from at least three sources: 11) passive tension exerted by the wing membrane may cause hindlimb motion; 22) hip and knee muscles may directly power hindlimb movement; and/or 33) vertical body oscillations could impose oscillations on the hindlimb via inertial effects

  • We found no effect of speed, individual, or their interaction on the flexion/ extension amplitude of ankle motion (ANOVA; speed: F1,14 = 1.859; p = 0.194; individual: F4,14 = 0.732; p = 0.211; interaction-term: F4,14 = 0.645; p = 0.640) or on the abduction/adduction amplitude of ankle motion

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Summary

Introduction

Bats are well known for their forelimbs, which are significantly modified into wings, but their hindlimbs are modified as well. Hindlimb movements contribute to three-dimensional wing shape. Potential aerodynamic consequences of wing shape alteration by these movements may include modulation of lift, drag, and pitching moment. The extent of such motion during bat flight is rarely documented, and it is not known whether hindlimb movements during flight are actively controlled or result from external forces acting on the limb. The bat hindlimb is rotated 90u or more relative to the ancestral mammal condition (Fig. 1). The knee is oriented dorsolaterally and its flexion moves the ankle and foot ventrally, rather than dorsally, the basal condition for mammals (Fig. 2). Knee flexion pulls the trailing edge of the wing ventrally rather than dorsally

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