[5] Many primate and bear species will adopt a bipedal gait in order to reach food or explore their environment, though there are a few cases where they walk on their hind limbs only.
Some animals commonly stand on their hind legs to reach food, keep watch, threaten a competitor or predator, or pose in courtship, but do not move bipedally.
While upright, non-locomotory limbs become free for other uses, including manipulation (in primates and rodents), flight (in birds), digging (in the giant pangolin), combat (in bears, great apes and the large monitor lizard) or camouflage.
[11] Even if one ignores exceptions caused by some kind of injury or illness, there are many unclear cases, including the fact that "normal" humans can crawl on hands and knees.
[18] 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.
Humans are the only primates who are normally biped, due to an extra curve in the spine which stabilizes the upright position, as well as shorter arms relative to the legs than is the case for the nonhuman great apes.
One hypothesis for human bipedalism is that it evolved as a result of differentially successful survival from carrying food to share with group members,[30] although there are alternative hypotheses.
[41] Recent evidence regarding modern human sexual dimorphism (physical differences between male and female) in the lumbar spine has been seen in pre-modern primates such as Australopithecus africanus.
[45] It has also been proposed that, like some modern-day apes, early hominins had undergone a knuckle-walking stage prior to adapting the back limbs for bipedality while retaining forearms capable of grasping.
[46] Numerous causes for the evolution of human bipedalism involve freeing the hands for carrying and using tools, sexual dimorphism in provisioning, changes in climate and environment (from jungle to savanna) that favored a more elevated eye-position, and to reduce the amount of skin exposed to the tropical sun.
In addition to the advantages of accruing from ability to carry objects – food or otherwise – the improvement of the visual range and the freeing of the hands for purposes of defence and offence may equally have played their part as catalysts.
[54] The evolution of an orthograde posture would have been very helpful on a savanna as it would allow the ability to look over tall grasses in order to watch out for predators, or terrestrially hunt and sneak up on prey.
[56] In fact, Elizabeth Vrba's turnover pulse hypothesis supports the savanna-based theory by explaining the shrinking of forested areas due to global warming and cooling, which forced animals out into the open grasslands and caused the need for hominids to acquire bipedality.
Findings also could shed light on discrepancies observed in the anatomy of A. afarensis, such as the ankle joint, which allowed it to "wobble" and long, highly flexible forelimbs.
If bipedalism started from upright navigation in trees, it could explain both increased flexibility in the ankle as well as long forelimbs which grab hold of branches.
In the face of long inter-birth intervals and low reproductive rates typical of the apes, early hominids engaged in pair-bonding that enabled greater parental effort directed towards rearing offspring.
Modern monogamous primates such as gibbons tend to be also territorial, but fossil evidence indicates that Australopithecus afarensis lived in large groups.
Early hominines such as Ardipithecus ramidus may have possessed an arboreal type of bipedalism that later independently evolved towards knuckle-walking in chimpanzees and gorillas[81] and towards efficient walking and running in modern humans (see figure).
Joseph Jordania from the University of Melbourne recently (2011) suggested that bipedalism was one of the central elements of the general defense strategy of early hominids, based on aposematism, or warning display and intimidation of potential predators and competitors with exaggerated visual and audio signals.
[82] Slow locomotion and strong body odor (both characteristic for hominids and humans) are other features often employed by aposematic species to advertise their non-profitability for potential predators.
A feedback mechanism from the advantages of bipedality in hot and open habitats would then in turn make a forest preadaptation solidify as a permanent state.
[87] This stone-tools theory is very unlikely, as though ancient humans were known to hunt, the discovery of tools was not discovered for thousands of years after the origin of bipedalism, chronologically precluding it from being a driving force of evolution.
"[89] It was then promoted by Elaine Morgan, as part of the aquatic ape hypothesis, who cited bipedalism among a cluster of other human traits unique among primates, including voluntary control of breathing, hairlessness and subcutaneous fat.
Since 2000 Carsten Niemitz has published a series of papers and a book[92] on a variant of the wading hypothesis, which he calls the "amphibian generalist theory" (German: Amphibische Generalistentheorie).
[96] The consequences of these two changes in particular resulted in painful and difficult labor due to the increased favor of a narrow pelvis for bipedalism being countered by larger heads passing through the constricted birth canal.
This is due to the fact that the narrowing of the hips and the change in the pelvic angle caused a discrepancy in the ratio of the size of the head to the birth canal.
The difficulties associated with simple standing in upright humans are highlighted by the greatly increased risk of falling present in the elderly, even with minimal reductions in control system effectiveness.
[101] In humans, walking is composed of several separate processes:[100] Early hominins underwent post-cranial changes in order to better adapt to bipedality, especially running.
As previously mentioned, longer hindlimbs assist in thermoregulation by reducing the total surface area exposed to direct sunlight while simultaneously allowing for more space for cooling winds.
In addition to the leg muscles, the increased size of the gluteus maximus in humans is an important adaptation as it provides support and stability to the trunk and lessens the amount of stress on the joints when running.