Bipedalism

Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs, or legs. An animal that usually moves in a bipedal manner is known as a biped, meaning "two feet" (from the Latin]] bi for "two" and ped for "foot"). Types of bipedal movement include walking, running, or hopping, on two appendages (typically legs).

Bipedalism evolved more than once in archosaurs, the group that includes both dinosaurs and crocodilians. A few lizards can also move bipedally, but only in emergencies. There are no known living or fossil bipedal amphibians.

Dinosaurs and their descendants

All dinosaurs are believed to be descended from a fully bipedal ancestor, perhaps similar to Eoraptor. It is believed that maniraptors were able to reach speeds of up to 65 km/h moving bipedally, comparable to their large, flightless-bird descendants such as the ostrich. Bipedal movement also re-evolved in a number of other dinosaur lineages such as the iguanodons. Some extinct members of the crocodilian line, a sister group to the dinosaurs and birds, also evolved bipedal forms - a crocodile relative from the triassic, Effigia okeeffeae, was believed to be bipedal.

Evolution

Recent evidence regarding modern human sexual dimorphism (physical differences between men and women) in the lumbar spine has been seen in pre-modern primates such as Australopithecus africanus. This dimorphism has been seen as an evolutionary adaptation of females to better bear lumbar load during pregnancy, an adaptation that non-bipedal primates would not need to make.

Bipedalism has a number of adaptive advantages, and has evolved independently in a number of lineages.

Early reptiles and lizards

The first known biped is the bolosaurid Eudibamus whose fossils date from 290 million years ago. Its long hindlegs, short forelegs, and distinctive joints all suggest bipedalism. This may have given increased speed. The species was extinct before the dinosaurs appeared.

Independent of Eudibamus, some modern lizard species have developed the capacity to run on their hind legs for added speed.

Dinosaurs and birds

Bipedalism also developed independently among the dinosaurs. Dinosaurs diverged from their archosaur ancestors approximately 230 million years ago during the Middle to Late Triassic period, roughly 20 million years after the Permian-Triassic extinction event wiped out an estimated 95% of all life on Earth. Radiometric dating of fossils from the early dinosaur genus Eoraptor establishes its presence in the fossil record at this time. Paleontologists believe Eoraptor resembles the common ancestor of all dinosaurs; if this is true, its traits suggest that the first dinosaurs were small, bipedal predators. The discovery of primitive, dinosaur-like ornithodirans such as Marasuchus and Lagerpeton in Argentinian Middle Triassic strata supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal predators.

Mammals (excluding humans)

A number of mammals will adopt a bipedal stance in specific situations such as for feeding or fighting. A number of groups of extant mammals have independently evolved bipedalism as their main form of locomotion - for example humans, giant pangolins, and macropods. Humans, as their bipedalism has been extensively studied are documented in the next section. Macropods are believed to have evolved bipedal hopping only once in their evolution, at some time no later than 45 million years ago.

Humans

There are at least twelve distinct hypotheses as to how and why bipedalism evolved in humans, and also some debate as to when. Bipedalism evolved well before the large human brain or the development of stone tools. The different hypotheses are not necessarily mutually exclusive and a number of selective forces may have acted together to lead to human bipedalism. It is important to distinguish between adaptations for bipedalism and adaptations for running, which came later still.

Various reasons have been proposed for the evolution of human bipedalism, including freeing the hands for tool use and carrying, sexual dimorphism in food gathering, changes in climate and habitat (from jungle to savanna) and to reduce the amount of skin exposed to the tropical sun. The first two explanations have been criticized for projecting modern social concerns and prejudices onto ancestral species. The latter two have been criticized for not making sense in the context of the forest and woodland biomes occupied by human ancestors. An alternative explanation is the mixture of savanna and scattered forests forced proto-humans to travel between clusters of trees and bipedalism offered greater efficiency for slow, long-distance travel between these clusters than knuckle-walking quadrupedism. However the wading hypothesis is increasingly gaining credence.