The functional morphology of hooding in cobras.

Many snakes, particularly cobras, form as part of a defensive display, a hood, an active lateral expansion of their neck skin and underlying musculature and ribs. We identified muscle groups possibly involved in hooding based on their attachments on the specialized ribs of the neck. We then used a combination of morphology, kinematic analysis, morphometrics, electromyography and muscle stimulation to test hypotheses about the functional basis of hooding. We confirmed that hood protraction and erection is an active process that begins cranially and extends caudally, often in stages, through the combined action of several sets of muscles. One set of axial muscles (levator costae and supracostalis lateralis superior) coursing along a line of action to rib displacement are the prime erectors acting to lift the hood. However, a second set of muscles connecting ribs to skin primarily keep the skin taut, rather than to displace the ribs relative to the vertebrae. A third set of muscles coursing between ribs function primarily to transmit forces between adjacent ribs rather than to move ribs. The maintenance of the erect hood requires continued muscle activity. Hood relaxation is due to both active muscle contraction of a fourth set of axial muscles and to passive recoil events in the costovertebral ligaments. The shape of the fully erect hood is reflective of the morphometrics of the underlying ribs, while the duration and kinematics of hood erection and relaxation are related to the behavioral context of the display.