Friday, March 13, 2009

Locomotion







The lack of limbs does not impede the movement of snakes, and they have developed several different modes of locomotion to deal with particular environments. Unlike the gaits of limbed animals, which form a continuum, each mode of snake locomotion is discrete and distinct from the others, and transitions between modes are abrupt.







Lateral undulation




Lateral undulation is the sole mode of aquatic locomotion, and the most common mode of terrestrial locomotion. In this mode, the body of the snake alternately flexes to the left and right, resulting in a series of rearward-moving 'waves'. While this movement appears rapid, snakes have been documented moving faster than two body-lengths per second, often much less.This mode of movement is similar to running in lizards of the same mass







Terrestrial




Terrestrial lateral undulation is the most common mode of terrestrial locomotion for most snake species.In this mode, the posteriorly moving waves push against contact points in the environment, such as rocks, twigs, irregularities in the soil, etc. Each of these environmental objects, in turn, generates a reaction force directed forward and towards the midline of the snake, resulting in forward thrust while the lateral components cancel out. The speed of this movement depends upon the density of push-points in the environment, with a medium density of about 8 along the snake's length being ideal. The wave speed is precisely the same as the snake speed, and as a result, every point on the snake's body follows the path of the point ahead of it, allowing snakes to move through very dense vegetation and small openings.







Aquatic




When swimming, the waves become larger as they move down the snake's body, and the wave travels backwards faster than the snake moves forwards. Thrust is generated by pushing their body against the water, resulting in the observed slip. In spite of overall similarities, studies show that the pattern of muscle activation is different in aquatic vs terrestrial lateral undulation, which justifies calling them separate modes. All snakes can laterally undulate forward (with backward-moving waves), but only sea snakes have been observed reversing the pattern, i.e. moving backwards via forward-traveling waves.







Sidewinding




Most often employed by colubroid snakes (colubrids, elapids, and vipers) when the snake must move in an environment which lacks any irregularities to push against (and which therefore renders lateral undulation impossible), such as a slick mud flat, or a sand dune. Sidewinding is a modified form of lateral undulation in which all of the body segments oriented in one direction remain in contact with the ground, while the other segments are lifted up, resulting in a peculiar 'rolling' motion. This mode of locomotion overcomes the slippery nature of sand or mud by pushing off with only static portions on the body, thereby minimizing slipping. The static nature of the contact points can be shown from the tracks of a sidewinding snake, which show each belly scale imprint, without any smearing. This mode of locomotion has very low caloric cost, less than ⅓ of the cost for a lizard or snake to move the same distance. Contrary to popular beliefs, there is no evidence that sidewinding is associated with hot sand




Concertina locomotion




When push-points are absent, but there is not enough space to use sidewinding because of lateral constraints, such as in tunnels, snakes rely on concertina locomotion. In this mode, the snake braces the posterior portion of its body against the tunnel wall while the front of the snake extends and straightens. The front portion then flexes and forms an anchor point, and the posterior is straightened and pulled forwards. This mode of locomotion is slow and very demanding, up to seven times the cost of laterally undulating over the same distance. This high cost is due to the repeated stops and starts of portions of the body as well as the necessity of using active muscular effort to brace against the tunnel walls.







Rectilinear locomotion




The slowest mode of snake locomotion is rectilinear locomotion, which is also the only one in which the snake does not need to bend its body laterally, though it may do so when turning. In this mode, the belly scales are lifted and pulled forward before being placed down and the body pulled over them. Waves of movement and stasis pass posteriorly, resulting in a series of ripples in the skin. The ribs of the snake do not move in this mode of locomotion and this method is most often used by large pythons, boas, and vipers when stalking prey across open ground as the snake's movements are subtle and harder to detect by their prey in this manner.







Other




The movement of snakes in arboreal habitats has only recently been studied. While on tree branches, snakes use several modes of locomotion depending on species and bark texture. In general, snakes will use a modified form of concertina locomotion on smooth branches, but will laterally undulate if contact points are available. Snakes move faster on small branches and when contact ponts are present, in contrast to limbed animals, which do better on large branches with little 'clutter'
Gliding snakes (Chrysopelea) of Southeast Asia launch themselves from branch tips, spreading their ribs and laterally undulating as they glide between trees. These snakes can perform a controlled glide for hundreds of feet depending upon launch altitude and can even turn in mid-air.

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