Bipedalism - Knowledge

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diameter of the branch, which can increase an arboreal feeding range and can be attributed to a convergent evolution of bipedalism evolving in arboreal environments. Hominine fossils found in dry grassland environments led anthropologists to believe hominines lived, slept, walked upright, and died only in those environments because no hominine fossils were found in forested areas. However, fossilization is a rare occurrence—the conditions must be just right in order for an organism that dies to become fossilized for somebody to find later, which is also a rare occurrence. The fact that no hominine fossils were found in forests does not ultimately lead to the conclusion that no hominines ever died there. The convenience of the savanna-based theory caused this point to be overlooked for over a hundred years.

1981:, holding onto supporting branches in order to navigate branches that were too flexible or unstable otherwise. In more than 75 percent of observations, the orangutans used their forelimbs to stabilize themselves while navigating thinner branches. Increased fragmentation of forests where A. afarensis as well as other ancestors of modern humans and other apes resided could have contributed to this increase of bipedalism in order to navigate the diminishing forests. 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. 2184:(1961) suggested that the carrying of meat "over considerable distances" (Hewes 1961:689) was the key factor. Isaac (1978) and Sinclair et al. (1986) offered modifications of this idea, as indeed did Lovejoy (1981) with his "provisioning model" described above. Others, such as Nancy Tanner (1981), have suggested that infant carrying was key, while others again have suggested stone tools and weapons drove the change. 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. (Wooden tools and spears fossilize poorly and therefore it is difficult to make a judgment about their potential usage.) 27: 1806:, an adaptation that non-bipedal primates would not need to make. Adapting bipedalism would have required less shoulder stability, which allowed the shoulder and other limbs to become more independent of each other and adapt for specific suspensory behaviors. In addition to the change in shoulder stability, changing locomotion would have increased the demand for shoulder mobility, which would have propelled the evolution of bipedalism forward. 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. 1830:) that favored a more elevated eye-position, and to reduce the amount of skin exposed to the tropical sun. It is possible that bipedalism provided a variety of benefits to the hominin species, and scientists have suggested multiple reasons for evolution of human bipedalism. There is also not only the question of why the earliest hominins were partially bipedal but also why hominins became more bipedal over time. For example, the postural feeding hypothesis describes how the earliest hominins became bipedal for the benefit of reaching food in trees while the savanna-based theory describes how the late hominins that started to settle on the ground became increasingly bipedal. 2434: 1954:. This hypothesis asserts that chimpanzees were only bipedal when they eat. While on the ground, they would reach up for fruit hanging from small trees and while in trees, bipedalism was used to reach up to grab for an overhead branch. These bipedal movements may have evolved into regular habits because they were so convenient in obtaining food. Also, Hunt's hypotheses states that these movements coevolved with chimpanzee arm-hanging, as this movement was very effective and efficient in harvesting food. When analyzing fossil anatomy, 2138:

groups of apes beginning to adopt bipedal postures more often. Others (e.g. Dart 1925) have offered the idea that the need for more vigilance against predators could have provided the initial motivation. Dawkins (e.g. 2004) has argued that it could have begun as a kind of fashion that just caught on and then escalated through sexual selection. And it has even been suggested (e.g. Tanner 1981:165) that male phallic display could have been the initial incentive, as well as increased sexual signaling in upright female posture.

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in a reduction in heat gain and helps heat dissipation. When a hominid is higher above the ground, the organism accesses more favorable wind speeds and temperatures. During heat seasons, greater wind flow results in a higher heat loss, which makes the organism more comfortable. Also, Wheeler explains that a vertical posture minimizes the direct exposure to the sun whereas quadrupedalism exposes more of the body to direct exposure. Analysis and interpretations of

7990: 8014: 8042: 8002: 601: 1933:, has a divergent big toe as well as the ankle strength to walk upright. "Little Foot" could grasp things using his feet like an ape, perhaps tree branches, and he was bipedal. Ancient pollen found in the soil in the locations in which these fossils were found suggest that the area used to be much more wet and covered in thick vegetation and has only recently become the arid desert it is now. 2370: 2449:

3:1, 2:1, 1:1, 5:2, and 3:2), although a 2:1 coupling ratio appears to be favored. Even though the evolution of bipedal gait has reduced the mechanical constraints on respiration in man, thereby permitting greater flexibility in breathing pattern, it has seemingly not eliminated the need for the synchronization of respiration and body motion during sustained running."

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metabolic rate via oxygen consumption, it was found that the quadrupedal and bipedal energy costs were very similar, implying that this transition in early ape-like ancestors would not have been very difficult or energetically costing. This increased travel efficiency is likely to have been selected for as it assisted foraging across widely dispersed resources.

298:— very rare genetic neurological disorders rather than normal behavior. 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. This article therefore avoids the terms "facultative" and "obligate", and focuses on the range of styles of locomotion 1844:

and reproductive success. Ko (2015) stated that there are two questions main regarding bipedalism 1. Why were the earliest hominins partially bipedal? and 2. Why did hominins become more bipedal over time? He argued that these questions can be answered with combination of prominent theories such as Savanna-based, Postural feeding, and Provisioning.

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the species became more bipedal, specialized feet would prevent the infant from conveniently clinging to the mother - hampering the mother's freedom and thus make her and her offspring more dependent on resources collected by others. Modern monogamous primates such as gibbons tend to be also territorial, but fossil evidence indicates that

2129:, threatening synchronous body movements, loud voice and extremely loud rhythmic singing/stomping/drumming on external subjects. 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. 208:, exclusively walk on two legs during the brief periods they spend on the ground. Many animals rear up on their hind legs while fighting or copulating. 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. 2192:

The observation that large primates, including especially the great apes, that predominantly move quadrupedally on dry land, tend to switch to bipedal locomotion in waist deep water, has led to the idea that the origin of human bipedalism may have been influenced by waterside environments. This idea,

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The thermoregulatory model explaining the origin of bipedalism is one of the simplest theories so far advanced, but it is a viable explanation. Dr. Peter Wheeler, a professor of evolutionary biology, proposes that bipedalism raises the amount of body surface area higher above the ground which results

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An alternative explanation is that the mixture of savanna and scattered forests increased terrestrial travel by proto-humans between clusters of trees, and bipedalism offered greater efficiency for long-distance travel between these clusters than quadrupedalism. In an experiment monitoring chimpanzee

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before losing it in Australopithecus. 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. It was also suggested in P. E. Wheeler's "The

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fossils from 4.2 to 3.9 million years ago and recent studies have suggested that obligate bipedal hominid species were present as early as 7 million years ago. Nonetheless, the evolution of bipedalism was accompanied by significant evolutions in the spine including the forward movement in position of

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engaged in pair-bonding that enabled greater parental effort directed towards rearing offspring. Lovejoy proposes that male provisioning of food would improve the offspring survivorship and increase the pair's reproductive rate. Thus the male would leave his mate and offspring to search for food and

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The form and function of modern-day humans' upper bodies appear to have evolved from living in a more forested setting. Living in this kind of environment would have made it so that being able to travel arboreally would have been advantageous at the time. Although different to human walking, bipedal

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Limited and exclusive bipedalism can offer a species several advantages. Bipedalism raises the head; this allows a greater field of vision with improved detection of distant dangers or resources, access to deeper water for wading animals and allows the animals to reach higher food sources with their

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while moving than do bipedal humans. "Quadrupedal species normally synchronize the locomotor and respiratory cycles at a constant ratio of 1:1 (strides per breath) in both the trot and gallop. Human runners differ from quadrupeds in that while running they employ several phase-locked patterns (4:1,

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Prehistoric fossil records show that early hominins first developed bipedalism before being followed by an increase in brain size. 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

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There are a variety of ideas which promote a specific change in behaviour as the key driver for the evolution of hominid bipedalism. For example, Wescott (1967) and later Jablonski & Chaplin (1993) suggest that bipedal threat displays could have been the transitional behaviour which led to some

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lived in large groups. However, while both gibbons and hominids have reduced canine sexual dimorphism, female gibbons enlarge ('masculinize') their canines so they can actively share in the defense of their home territory. Instead, the reduction of the male hominid canine is consistent with reduced

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males were nearly twice the weight of females. However, Lovejoy's model posits that the larger range a provisioning male would have to cover (to avoid competing with the female for resources she could attain herself) would select for increased male body size to limit predation risk. Furthermore, as

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Sigmon (1971) demonstrated that chimpanzees exhibit bipedalism in different contexts, and one single factor should be used to explain bipedalism: preadaptation for human bipedalism. Day (1986) emphasized three major pressures that drove evolution of bipedalism: food acquisition, predator avoidance,

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Shoulder stability would decrease with the evolution of bipedalism. Shoulder mobility would increase because the need for a stable shoulder is only present in arboreal habitats. Shoulder mobility would support suspensory locomotion behaviors which are present in human bipedalism. The forelimbs are

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Respiration through bipedality means that there is better breath control in bipeds, which can be associated with brain growth. The modern brain utilizes approximately 20% of energy input gained through breathing and eating, as opposed to species like chimpanzees who use up twice as much energy as

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by reducing the total surface area exposed to direct sunlight while simultaneously allowing for more space for cooling winds. Additionally, having longer limbs is more energy-efficient, since longer limbs mean that overall muscle strain is lessened. Better energy efficiency, in turn, means higher

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Charles Darwin wrote that "Man could not have attained his present dominant position in the world without the use of his hands, which are so admirably adapted to the act of obedience of his will". Darwin (1871:52) and many models on bipedal origins are based on this line of thought. Gordon Hewes

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came down from the tree's branches and adapted to life on the savanna by walking erect on two feet. The theory suggests that early hominids were forced to adapt to bipedal locomotion on the open savanna after they left the trees. One of the proposed mechanisms was the knuckle-walking hypothesis,

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It seems unlikely that any single factor was responsible for such a dramatic change in behaviour. 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

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Other theories have been proposed that suggest wading and the exploitation of aquatic food sources (providing essential nutrients for human brain evolution or critical fallback foods) may have exerted evolutionary pressures on human ancestors promoting adaptations which later assisted full-time

2216:", as originally formulated, has not been accepted or considered a serious theory within the anthropological scholarly community. Others, however, have sought to promote wading as a factor in the origin of human bipedalism without referring to further ("aquatic ape" related) factors. Since 2000 2124:

of potential predators and competitors with exaggerated visual and audio signals. According to this model, hominids were trying to stay as visible and as loud as possible all the time. Several morphological and behavioral developments were employed to achieve this goal: upright bipedal posture,

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Most bipedal animals move with their backs close to horizontal, using a long tail to balance the weight of their bodies. The primate version of bipedalism is unusual because the back is close to upright (completely upright in humans), and the tail may be absent entirely. Many primates can stand

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can be used to quantify the whole-body kinetic & potential energy, with walking displaying an out-of-phase relationship indicating exchange between the two. This model applies to all walking organisms regardless of the number of legs, and thus bipedal locomotion does not differ in terms of

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upright. It is possible that bipedalism evolved in the trees, and was later applied to the savanna as a vestigial trait. Humans and orangutans are both unique to a bipedal reactive adaptation when climbing on thin branches, in which they have increased hip and knee extension in relation to the

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species move bipedally when running, usually to escape from threats. 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. Several arboreal primate species, such as

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other than infants normally walk and run in biped fashion, but almost all can crawl on hands and knees when necessary. There are even reports of humans who normally walk on all fours with their feet but not their knees on the ground, but these cases are a result of conditions such as

2510:. Recently, spurred by the success of creating a fully passive, un-powered bipedal walking robot, those working on such machines have begun using principles gleaned from the study of human and animal locomotion, which often relies on passive mechanisms to minimize power consumption. 2257:

appears to be rare and actively avoided cross-culturally, even if birthing methods may differ between said cultures. 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

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Early hominins underwent post-cranial changes in order to better adapt to bipedality, especially running. One of these changes is having longer hindlimbs proportional to the forelimbs and their effects. As previously mentioned, longer hindlimbs assist in

640:, although usually quadrupedal, will sometimes move between adjacent feeding patches with a squatting, shuffling bipedal form of locomotion. However, they can only do so for brief amounts, as their bodies are not adapted for constant bipedal locomotion. 1776:

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. Bipedal specializations are found in

2457:. This is because breath control means that the muscles associated with breathing can be manipulated into creating sounds. This means that the onset of bipedality, leading to more efficient breathing, may be related to the origin of verbal language. 2203:

who said: "It seems to me likely that Man learnt to stand erect first in water and then, as his balance improved, he found he became better equipped for standing up on the shore when he came out, and indeed also for running." It was then promoted by

2175:, rather than the hotter conditions being hypothetically bipedalism's initial stimulus. 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. 2096:

may have possessed an arboreal type of bipedalism that later independently evolved towards knuckle-walking in chimpanzees and gorillas and towards efficient walking and running in modern humans (see figure). It is also proposed that one cause of

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evolution of bipedality and loss of functional body hair in hominids", that a possible advantage of bipedalism in the savanna was reducing the amount of surface area of the body exposed to the sun, helping regulate body temperature. In fact,

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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. The evolution of

549:. 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. 2069:

suggest that the species was closely related to African-ape ancestors. This possibly provides a species close to the true connection between fully bipedal hominins and quadruped apes. According to Richard Dawkins in his book

2022:, suggests a reduction in inter-male antagonistic behavior in early hominids. In addition, this model is supported by a number of modern human traits associated with concealed ovulation (permanently enlarged breasts, lack of 1900:

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.

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Others state hominines had already achieved the bipedal adaptation that was used in the savanna. The fossil evidence reveals that early bipedal hominins were still adapted to climbing trees at the time they were also

4974: 2287:. 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. 288:

Zoologists often label behaviors, including bipedalism, as "facultative" (i.e. optional) or "obligate" (the animal has no reasonable alternative). Even this distinction is not completely clear-cut — for example,

2086:. These apes may have once been bipedal, but then lost this ability when they were forced back into an arboreal habitat, presumably by those australopithecines from whom eventually evolved hominins. Early 2661:

Heglund, NC; Cavagna, GA; Taylor, CR (1982). "Energetics and mechanics of terrestrial locomotion. III. Energy changes of the centre of mass as a function of speed and body size in birds and mammals".

2222: 663:. One hypothesis for human bipedalism is that it evolved as a result of differentially successful survival from carrying food to share with group members, although there are alternative hypotheses. 2421:

muscles of the thigh are both so crucial to bipedal activities that each alone is much larger than the well-developed biceps of the arms. In addition to the leg muscles, the increased size of the

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Running is characterized by a spring-mass movement. Kinetic and potential energy are in phase, and the energy is stored & released from a spring-like limb during foot contact, achieved by the

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However, this model has been debated, as others have argued that early bipedal hominids were instead polygynous. Among most monogamous primates, males and females are about the same size. That is

1923:, found in Hadar in Ethiopia, which may have been forested at the time of Lucy's death, had curved fingers that would still give her the ability to grasp tree branches, but she walked bipedally. " 682:

and Poko (chimpanzee), were found to move bipedally. Natasha switched to exclusive bipedalism after an illness, while Poko was discovered in captivity in a tall, narrow cage. Oliver reverted to

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used by various groups of animals. Normal humans may be considered "obligate" bipeds because the alternatives are very uncomfortable and usually only resorted to when walking is impossible.

2574: 588:, became famous locally and on the internet for having a frequent bipedal gait, although this is attributed to injuries on the bear's front paws. A two-legged fox was filmed in a 3205:

Sereno, Paul C.; Catherine A. Forster; Raymond R. Rogers; Alfredo M. Monetta (January 1993). "Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria".

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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

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can exceed 100 km/h (62 mph). Even though bipedalism is slower at first, over long distances, it has allowed humans to outrun most other animals according to the

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will fight in a bipedal stance to use their forelegs as weapons. A number of mammals will adopt a bipedal stance in specific situations such as for feeding or fighting.

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Isbell LA, Young TP (1996). "The evolution of bipedalism in hominids and reduced group size in chimpanzees: alternative responses to decreasing resource availability".

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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

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return carrying the food in his arms walking on his legs. This model is supported by the reduction ("feminization") of the male canine teeth in early hominids such as

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bipedalism. It has also been thought that consistent water-based food sources had developed early hominid dependency and facilitated dispersal along seas and rivers.

2212:, who cited bipedalism among a cluster of other human traits unique among primates, including voluntary control of breathing, hairlessness and subcutaneous fat. The " 387:

whose fossils date from 290 million years ago. Its long hind-legs, short forelegs, and distinctive joints all suggest bipedalism. The species became extinct in the

195:) as well as various other extinct groups evolving the trait independently. A larger number of modern species intermittently or briefly use a bipedal gait. Several 814:

walk bipedally on the sea floor using two of their arms, allowing the remaining arms to be used to camouflage the octopus as a mat of algae or a floating coconut.

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The great majority of living terrestrial vertebrates are quadrupeds, with bipedalism exhibited by only a handful of living groups. Humans, gibbons and large birds

6703:, "The Archeological Evidence for the Activities of Early African Hominids" In:Early Hominids of Africa (Jolly, C.J. (Ed.)), Duckworth (London), 219–254, (1978). 2494:

involved only wheels, treads, or multiple legs. Recent cheap and compact computing power has made two-legged robots more feasible. Some notable biped robots are

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Crompton, R. H.; Thorpe, S. K. S. (2007-11-16). "Response to Comment on "Origin of Human Bipedalism As an Adaptation for Locomotion on Flexible Branches"".

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will squat on their hindlegs to manipulate some objects but revert to four limbs when moving (the beaver will move bipedally if transporting wood for their

2026:) and low sperm competition (moderate sized testes, low sperm mid-piece volume) that argues against recent adaptation to a polygynous reproductive system. 4144: 3670: 3014: 348:

by raising one foot at a time. On the other hand, most macropods, smaller birds, lemurs and bipedal rodents move by hopping on both legs simultaneously.

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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.

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Green, Alemseged, David, Zeresenay (2012). "Australopithecus afarensis Scapular Ontogeny, Function, and the Role of Climbing in Human Evolution".

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locomotion in trees was thought to be advantageous. It has also been proposed that, like some modern-day apes, early hominins had undergone a

5746: 3486: 3398:"Locomotor versatility in the white-handed gibbon (Hylobates lar): A spatiotemporal analysis of the bipedal, tripedal, and quadrupedal gaits" 511:, a sister group to the avemetatarsalians (the group including dinosaurs and relatives), also evolved bipedal forms – a poposauroid from the 2989: 3156: 352:

are able to walk or hop, most commonly alternating feet when moving arboreally and hopping on both feet simultaneously when on the ground.

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Bipedal movement occurs in a number of ways and requires many mechanical and neurological adaptations. Some of these are described below.

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hip and hind limb very clearly indicate bipedalism, but these fossils also indicate very inefficient locomotive movement when compared to

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Thorpe, S. K.; Holder, R. L.; Crompton, R. H. (2007). "Origin of human bipedalism as an adaptation for locomotion on flexible branches".

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after developing arthritis. Non-human primates often use bipedal locomotion when carrying food, or while moving through shallow water.

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of bipedalism was linked to monogamy. In the face of long inter-birth intervals and low reproductive rates typical of the apes, early

7123: 7083: 3877: 3444: 3282:"The Phylogenetic Position of the Musky Rat-Kangaroo and the Evolution of Bipedal Hopping in Kangaroos (Macropodidae: Diprotodontia)" 2220:

has published a series of papers and a book on a variant of the wading hypothesis, which he calls the "amphibian generalist theory" (

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which states that human ancestors used quadrupedal locomotion on the savanna, as evidenced by morphological characteristics found in

1467: 7975: 7819: 7133: 2262:. The result of this is that there is greater difficulty in birthing for hominins in general, let alone to be doing it by oneself. 1771: 1487: 856: 648: 560:. All primates possess some bipedal ability, though most species primarily use quadrupedal locomotion on land. Primates aside, the 533:

A number of groups of extant mammals have independently evolved bipedalism as their main form of locomotion - for example humans,

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move bipedally by hopping. Very few non-primate mammals commonly move bipedally with an alternating leg gait. Exceptions are the

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Non-human primates habitually deliver their young on their own, but the same cannot be said for modern-day humans. Isolated

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The maximum bipedal speed appears slower than the maximum speed of quadrupedal movement with a flexible backbone – both the

6830: 5686:"Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor" 2063:

and was reduced in chimpanzee and gorilla when they became more specialized. Other recent studies of the foot structure of

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strata supports this view; analysis of recovered fossils suggests that these animals were indeed small, bipedal predators.

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become bipedal during high-speed, sprint locomotion, including the world's fastest lizard, the spiny-tailed iguana (genus

7839: 6629: 5052: 3671:"Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): Testing hypotheses on the evolution of bipedalism" 2332:

humans have that non-human apes do not. Rather, walking is characterized by an "inverted pendulum" movement in which the

5928: 2328:, which requires the engagement of both the hip and the knee joints. This human ability to walk is made possible by the 1970:. For this reason, Hunt argues that bipedalism evolved more as a terrestrial feeding posture than as a walking posture. 7948: 6802: 6198: 4891:

Java man: how two geologists' dramatic discoveries changed our understanding of the evolutionary path to modern humans

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humans for the same amount of movement. This excess energy, leading to brain growth, also leads to the development of

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freed from weight-bearing requirements, which makes the shoulder a place of evidence for the evolution of bipedalism.

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Energy-efficient means of standing bipedally involve constant adjustment of balance, and of course these must avoid

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Jablonski, N.G.; Chaplin, G. (1993). "Origin of Habitual Terrestrial Bipedalism in the Ancestor of the Hominidae".

5010: 3323:"A new Pleistocene tree-kangaroo (Diprotodontia: Macropodidae) from the Nullarbor Plain of south-central Australia" 1427: 8074: 7958: 7854: 1637: 317:. Staying still on both legs. In most bipeds this is an active process, requiring constant adjustment of balance. 245:

become free for other uses, including manipulation (in primates and rodents), flight (in birds), digging (in the

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has very similar features of the hand and shoulder to the chimpanzee, which indicates hanging arms. Also, the

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Several groups of modern species are habitual bipeds whose normal method of locomotion is two-legged. In the

7076: 3259: 1990: 5361:"An enlarged postcranial sample confirms Australopithecus afarensis dimorphism was similar to modern humans" 3855:"Bipedal orangs, gait of a dinosaur, and new-look Ichthyostega: exciting times in functional anatomy part I" 3564:

Daver G, Guy F, Mackaye HT, Likius A, Boisserie J, Moussa A, Pallas L, Vignaud P, Clarisse ND (2022-08-24).

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Some of the fossils found actually showed that there was still an adaptation to arboreal life. For example,

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Napier (1963) argued that it is unlikely that a single factor drove the evolution of bipedalism. He stated "

7824: 7701: 7587: 6794: 5105:"Arboreal bipedalism in wild chimpanzees: Implications for the evolution of hominid posture and locomotion" 3254: 2584: 1802:. This dimorphism has been seen as an evolutionary adaptation of females to bear lumbar load better during 1077: 849: 4157:

This article has good pictures explaining the differences between bipedal and non-bipedal pregnancy loads.

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in the foot and leg, respectively. Again, the whole-body kinetics are similar to animals with more limbs.

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were previously thought to have been bipedal, but recent trackways have all shown quadrupedal locomotion.

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Pontzer, H.; Raichlen, D. A.; Rodman, P. S. (2014). "Bipedal and quadrupedal locomotion in chimpanzees".

826: 6387: 4990:"Walk Like an Orangutan: Ape's stroll through the trees may shed light on evolution of human bipedalism" 3039: 8032: 7780: 7711: 7247: 7240: 7233: 7050: 5866:"Bipedality and hair loss in human evolution revisited: The impact of altitude and activity scheduling" 5829:

Wheeler, P.E. (1991). "The influence of bipedalism on the energy and water budgets of early hominids".

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Richmond, B. G.; Strait, D. S. (2000). "Evidence that humans evolved from a knuckle-walking ancestor".

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by larger heads passing through the constricted birth canal. This phenomenon is commonly known as the

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Wheeler, P. E. (1990). "The influence of thermoregulatory selection pressures on hominid evolution".

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forelimbs, and that it is less parsimonious to assume that knuckle walking developed twice in genera

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has been hypothesized to improve locomotor performance, which could aid in escaping from predators.

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Wheeler, P. E. (1984). "The Evolution of Bipedality and Loss of Functional Body Hair in Hominoids".

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Wheeler, P. E. (1984) "The Evolution of Bipedality and Loss of Functional Body Hair in Hominoids."

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Wheeler, P. E. (1984). "The evolution of bipedality and loss of functional body hair in hominids".

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with bipedal sideways hopping movements of the hind legs, holding their forelimbs up for balance.

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Harmon E (2009). "Size and shape variation in the proximal femur of Australopithecus africanus".

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Davies, S.J.J.F. (2003). "Birds I Tinamous and Ratites to Hoatzins". In Hutchins, Michael (ed.).

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Huffard CL, Boneka F, Full RJ (2005). "Underwater bipedal locomotion by octopuses in disguise".

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Other mammals engage in limited, non-locomotory, bipedalism. A number of other animals, such as

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Bauer, Harold (1976). "Chimpanzee bipedal locomotion in the Gombe National Park, East Africa".

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DeSilva, J., "First Steps: How Upright Walking Made Us Human" HarperCollins (New York), (2021)

6314:"Spinopelvic pathways to bipedality: why no hominids ever relied on a bent-hip-bent-knee gait" 6091: 4025: 4017: 3512:"Primate Factsheets: Gelada baboon (Theropithecus gelada) Taxonomy, Morphology, & Ecology" 3511: 1950:

The postural feeding hypothesis has been recently supported by Dr. Kevin Hunt, a professor at

435:. All dinosaurs are thought to be descended from a fully bipedal ancestor, perhaps similar to 7311: 6914: 6034: 5308:

White TD, et al. (2009). "Ardipithecus ramidus and the paleobiology of early hominids".

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began in primates about four million years ago, or as early as seven million years ago with

7892: 7795: 7696: 7657: 7602: 7507: 7211: 7204: 6716: 6580: 6278: 5877: 5838: 5768: 5697: 5576: 5525: 5417: 5317: 5259: 5197: 5157: 5151: 4924: 4862: 4811: 4746: 4683: 4620: 4577: 4501: 4433: 4335: 4264: 4176: 4101: 3982: 3790: 3580: 3409: 3214: 3121: 3071: 2954: 2919: 2860: 2758: 2194: 2092: 2065: 2055: 2010: 1706: 1507: 1354: 1223: 1208: 794:. Many reptile species will also temporarily adopt bipedalism while fighting. One genus of 3881: 3371: 2910:

Djawdan, M (1993). "Locomotor performance of bipedal and quadrupedal heteromyid rodents".

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stage prior to adapting the back limbs for bipedality while retaining forearms capable of

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Sylvester, Adam D. (2006). "Locomotor Coupling and the Origin of Hominin Bipedalism".

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Sylvester, Adam D. (2006). "Locomotor Coupling and the Origin of Hominin Bipedalism".

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Sigmon, Becky (1971). "Bipedal behavior and the emergence of erect posture in man".

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Sigmon, Becky (1971). "Bipedal behavior and the emergence of erect posture in man".

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when threatened, rearing up on its front legs while facing the attacker so that its

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establishes its presence in the fossil record at this time. Paleontologists suspect

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will stand on hind legs to survey their surroundings, but will not walk bipedally.

675: 151:, all the early forms and many later groups were habitual or exclusive bipeds; the 75: 6752: 6735:

Tanner, N. M., "On Becoming Human", Cambridge University Press (Cambridge), (1981)

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Dean, F. 2000. Primate diversity. W.W. Norton & Company, Inc: New York. Print.

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the well muscled legs, against the small and bony wings. Likewise in humans, the

2395: 2226: 2217: 2171:. This then allowed for the more efficient exploitation of the hotter conditions 2117: 2109: 2023: 1811: 1322: 1091: 1047: 720: 683: 577: 534: 493: 323:. One foot in front of another, with at least one foot on the ground at any time. 2406:

Bipedalism requires strong leg muscles, particularly in the thighs. Contrast in

329:. One foot in front of another, with periods where both feet are off the ground. 8058: 7471: 7175: 7161: 7100: 7001: 6767: 6653: 6290: 4188: 3592: 3133: 2529: 2470: 2466: 2441: 2360:

Rotation of the hips about the horizontal axis to improve balance during stance

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are known to move bipedally at high speeds. Bipedalism is rarely found outside

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Brunet M, Guy F, Pilbeam D, Mackaye HT, Likius A, et al. (11 July 2002).

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suggest bipedalism. It is thus possible that bipedalism evolved very early in

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There are a number of states of movement commonly associated with bipedalism.

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Suwa G, Kono RT, Simpson SW, Asfaw B, Lovejoy CO, White TD (2 October 2009).

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antelope stands on its hind legs while eating from trees, as did the extinct

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Kuliukas, A. (2013). "Wading Hypotheses of the Origin of Human Bipedalism".

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One theory on the origin of bipedalism is the behavioral model presented by

7645: 7633: 7575: 7449: 7397: 7362: 7287: 7194: 7027: 6873: 6728: 6663: 6455: 6427: 6363: 6329: 6298: 6140: 5907: 5729: 5670: 5596: 5545: 5492: 5437: 5394: 5376: 5337: 5279: 5217: 5136: 4944: 4874: 4831: 4766: 4640: 4365: 4300: 4196: 4121: 3951: 3694: 3608: 3429: 3395: 3307: 3149:"Global Warming Led To Atmospheric Hydrogen Sulfide And Permian Extinction" 3091: 2888: 2744: 2718: 2549: 2391: 2357:

Rotation of the hips about the axis of the spine, to increase stride length

2325: 2148: 2121: 2082: 1880: 1793: 1738: 1312: 1279: 1193: 1117: 1104: 756: 748: 671:

Injured chimpanzees and bonobos have been capable of sustained bipedalism.

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Bipedalism: Pressures, origins and modes. Major topics in human evolution

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A short 'push' from the ankle prior to toe-off, propelling the swing leg

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bipedalism preceded further refinement of bipedalism by the pressure of

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ancestors approximately 230 million years ago during the Middle to Late

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Berman, David S.; et al. (2000). "Early Permian Bipedal Reptile".

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can run bipedally across the surface of water for some distance. Among

613: 589: 573: 485: 479: 432: 428: 388: 379: 366: 172: 8053: 6247: 6215: 5473: 1977:, Professor Susannah Thorpe examined the most arboreal great ape, the 712: 7581: 7486: 4632: 4090:"Fetal load and the evolution of lumbar lordosis in bipedal hominins" 3226: 2519: 2418: 2384: 2087: 2060: 1994: 1978: 1859: 1803: 1722: 1399: 803: 799: 776: 772: 736: 633: 557: 522: 490: 445: 412: 383: 335:/hopping. Moving by a series of jumps with both feet moving together. 205: 140: 136: 6686:

Hewes, G. W., "Food Transport and the Origin of Hominid Bipedalism"

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Das Geheimnis des Aufrechten Gangs ~ Unsere Evolution Verlief Anders

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reveal that this hypothesis needs modification to consider that the

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inter-male aggression in a pair-bonded though group living primate.

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Unlike non-human apes that are able to practice bipedality such as

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6117:"Shallow-water habitats as sources of fallback foods for hominins" 3774: 3538:

Primate morphophysiology, locomotor analyses, and human bipedalism

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labelled "the wading hypothesis", was originally suggested by the

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bipedalism is rare, but it is found in the "reared-up" running of

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Burk, Angela; Michael Westerman; Mark Springer (September 1988).

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Wrangham R, Cheney D, Seyfarth R, Sarmiento E (December 2009).

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afarensisassemblages confirm only moderate skeletal dimorphism"

5149: 3372:"Two-legged fox is nature conquering all, says wildlife expert" 2259: 2152: 1823: 783: 708: 637: 629: 625: 617: 553: 542: 201: 196: 160: 58: 5183:"A new hominid from the Upper Miocene of Chad, Central Africa" 4888:

Lewin, Roger; Swisher, Carl Celso; Curtis, Garniss H. (2000).

2344:

In humans, walking is composed of several separate processes:

2125:

longer legs, long tightly coiled hair on the top of the head,

419:

chicks have claws on their wings which they use for climbing.

7667: 7558: 7548: 3554: 2495: 2487: 2479: 2254: 1967: 1815: 461: 405: 290: 217: 155:

are members of a clade of exclusively bipedal dinosaurs, the

102: 6666:, "Australopithecus africanus: The Ape Man of South Africa" 5511:"Reexamining human origins in light of Ardipithecus ramidus" 2369: 7326: 6847: 3396:

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2959:"Maximal running speeds of bipedal and quadrupedal rodents" 2507: 2499: 1993:, known as "male provisioning". Lovejoy theorizes that the 763:, capable of spraying an offensive oil, face its attacker. 716: 163:, habitual bipedalism has evolved multiple times, with the 152: 90: 84: 5610:

Keith Oatley; Dacher Keltner; Jennifer M. Jenkins (2006).

1822:

in provisioning, changes in climate and environment (from

268:

can reach speeds of 70 km/h (43 mph), while the

49:

where an animal moves by means of its two rear (or lower)

6419:

A moving topic: control and dynamics of animal locomotion

6088:

Survival of the fattest: the key to human brain evolution

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Thorpe, S. K. S.; Holder, R. L.; Crompton, R. H. (2007).

3339:

10.1671/0272-4634(2008)28[463:ANPTDM]2.0.CO;2

1755: 728: 700: 250: 54: 4709: 4707: 4705: 4087: 2946: 2475: 456:

wiped out an estimated 95 percent of all life on Earth.

5180: 4088:

Whitcome KK, Shapiro LJ, Lieberman DE (December 2007).

3878:"The Story behind the Picture - Monitor Lizards Combat" 2844:

Bramble, Dennis M.; Lieberman, Daniel E. (2004-11-18).

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gracile type species while gorillas are descended from

592:

garden in 2023, most likely having been born that way.

5011:"Orangutans Show First Walking May Have Been on Trees" 3008:"Human Hand-Walkers: Five Siblings Who Never Stood Up" 2952: 2048: 1792:(physical differences between male and female) in the 499:

Bipedal movement also re-evolved in a number of other

283: 8030: 6161: 4914: 4702: 3563: 2437:

The human respiratory system, encased by the rib cage

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Lovejoy, C.O. (1988). "Evolution of Human walking".

3906:"Bipedal animals, and their differences from humans" 2660: 93: 87: 81: 6658:

The Descent of Man and Selection in Relation to Sex

5863: 3171:"The Day The Earth Nearly Died - programme summary" 821: 78: 7571:(archaic homo sapiens, anatomically modern humans) 4051:"Becoming Human: The Evolution of Walking Upright" 3638:"What Does It Mean To Be Human? - Walking Upright" 3321:Prideaux, Gavin J.; Warburton, Natalie M. (2008). 3250:"Dino-Era Fossil Reveals Two-Footed Croc Relative" 3175:Science & Nature > TV & Radio Follow-up 2486:For nearly the whole of the 20th century, bipedal 2033:is minimal, and other studies have suggested that 1936: 6693:Hunt, K. D., "The Evolution of Human Bipedality" 6370: 5074:"Walking the walk: evolution of human bipedalism" 4074:"A pregnant woman's spine is her flexible friend" 3929: 3320: 2701:Clemente, Christofer J.; Wu, Nicholas C. (2018). 2640:. Austin, TX: Raintree Steck-Vaughn. p. 22. 767:Limited bipedalism in non-mammals (and non-birds) 612:upright on their hind legs without any support. 8066: 6706: 6216:"The Evolution of Bipedalism and Assisted Birth" 6090:. 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Duke University Lemur Center. Archived from 1945: 7077: 6838: 6824: 5683: 5504: 5502: 3972: 3479: 2846:"Endurance running and the evolution of Homo" 1841:equally have played their part as catalysts." 1796:has been seen in pre-modern primates such as 850: 431:, the group that includes both dinosaurs and 373: 34:, the co-fastest extant biped along with the 6680:", Weidenfeld and Nicolson (London), (2004). 5747:Why do People Sing? Music in Human Evolution 4956: 4954: 4787:: CS1 maint: multiple names: authors list ( 4655: 4560:Brazilian Archives of Biology and Technology 4416:Brazilian Archives of Biology and Technology 4071: 3923: 3668: 3630: 3535: 2903: 2387:, particularly when running long distances. 564:(kangaroos, wallabies and their relatives), 7556: 6406: 6051: 5450: 4852: 3756:"Monkey apes humans by walking on two legs" 3107:"The evolution of locomotion in archosaurs" 2351:Passive ballistic movement of the swing leg 2132: 628:exhibit forms of bipedalism. On the ground 452:period, roughly 20 million years after the 112:'foot'). Types of bipedal movement include 7084: 7070: 6831: 6817: 5992: 5499: 5150:T. Douglas Price; Gary M. Feinman (2003). 5053:"Our upright walking started in the trees" 4557: 3146: 3104: 2700: 2566: 857: 843: 715:, as will the raccoon when holding food). 6445: 6353: 6268: 6213: 5927:. Cambridge: Cambridge University Press. 5897: 5719: 5709: 5660: 5642: 5482: 5472: 5384: 5109:American Journal of Physical Anthropology 5032:"Upright orangutans point way to walking" 4951: 4571: 4495: 4484:American Journal of Physical Anthropology 4427: 4381:American Journal of Physical Anthropology 4355: 4227: 4166: 4020:. In Michael Ruse; Joseph Travis (eds.). 3831:. University of Liverpool. Archived from 3675:American Journal of Physical Anthropology 3449:American Journal of Physical Anthropology 3297: 3273: 3191:Hayward, T. (1997). 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Tokyo: University of Tokyo Press. 3442: 2696: 2694: 2692: 1694: 1672: 1650: 1626: 1606: 1586: 1562: 1542: 1519: 1496: 1476: 1456: 1436: 1416: 824: 674:Three captive primates, one macaque 7091: 5966:"Was man more aquatic in the past?" 5156:. Boston, MA: McGraw Hill. p. 4981: 4542: 4145:"Why Pregnant Women Don't Tip Over" 2811:"Timed running speed of a cheetah ( 2049:Early bipedalism in homininae model 1833: 657:or about 12 million years ago with 521:, is thought to have been bipedal. 460:of fossils from the early dinosaur 422: 408:are bipeds, as is the case for all 284:Facultative and obligate bipedalism 13: 6803:National Museum of Natural History 6647: 6569:"Running and Breathing in Mammals" 5365:Philos Trans R Soc Lond B Biol Sci 4413: 3995:10.1038/scientificamerican1188-118 3908:. Ingentaconnect.com. 2004-05-01. 3327:Journal of Vertebrate Paleontology 3195:. Orbis Publishing Ltd. D36040612. 2831:10.1111/j.1469-7998.1997.tb04840.x 2782:10.1111/j.1469-7998.1983.tb02087.x 2611:Grzimek's Animal Life Encyclopedia 2444:, have more restrictive breathing 2178: 2105:Warning display (aposematic) model 355: 339: 147:) developed bipedalism; among the 14: 8096: 6783: 6389:Muscles, reflexes, and locomotion 6164:Trends in Ecology & Evolution 5625:Prang, Thomas Cody (2019-04-30). 4072:Steve Connor (13 December 2007). 2689: 2460: 1927:", a nearly-complete specimen of 810:, though at least two species of 454:Permian-Triassic extinction event 8052: 8040: 8012: 8000: 7989: 7988: 6614: 6560: 6305: 5931:from the original on 2013-05-22. 5921:Tanner, Nancy Makepeace (1981). 5297:from the original on 2022-10-09. 5092:from the original on 2022-10-09. 4977:from the original on 2017-03-05. 4967:South African Journal of Science 3880:. Wildlife Times. Archived from 3853:Naish, Darren (April 28, 2008). 3626:from the original on 2022-08-25. 3181:from the original on 2012-09-01. 3159:from the original on 2011-06-05. 3020:from the original on 2008-09-10. 2995:from the original on 2010-06-16. 2348:Vaulting over a stiff stance leg 2187: 771:Bipedalism is unknown among the 74: 6632:from the original on 2013-11-07 6422:. Vol. 6. pp. 387–8. 6262: 6207: 6182: 6155: 6108: 6079: 5954: 5935: 5914: 5857: 5822: 5787: 5752: 5736: 5677: 5618: 5603: 5560: 5444: 5401: 5352: 5301: 5232: 5174: 5153:Images of the Past, 5th edition 5096: 4908: 4881: 4846: 4795: 4730: 4667: 4604: 4551: 4536: 4475: 4471:. Archives de Biologie (Liege). 4460: 4407: 4372: 4315: 4244: 4203: 4160: 4136: 4065: 4049:Wayman, Erin (August 6, 2012). 4042: 4009: 3966: 3912:from the original on 2012-11-26 3898: 3869: 3846: 3817: 3747: 3712: 3662: 3518:from the original on 2012-05-09 3504: 3436: 3389: 3364: 3353:from the original on 2011-10-19 3314: 3241: 3198: 3185: 3163: 3140: 3098: 3053: 3024: 2663:Journal of Experimental Biology 2308:Profile view of the human spine 2273: 2240: 2231:Amphibische Generalistentheorie 1937:Traveling efficiency hypothesis 1879:instead of evolving it once as 38:, at 70 km/h (43 mph) 6392:. Princeton University Press. 6271:Journal of Theoretical Biology 6232:10.1525/maq.1996.10.2.02a00100 6220:Medical Anthropology Quarterly 5981:(174): 642–645. Archived from 4169:Journal of Theoretical Biology 3876:Sharma, Jayanth (2007-03-08). 3248:Handwerk, Brian (2006-01-26). 2837: 2654: 2542: 2428: 2401: 2101:was a less efficient running. 1415: 580:and in some circumstances the 541:, numerous species of jumping 444:Dinosaurs diverged from their 1: 6753:10.1525/aa.1967.69.6.02a00110 6193:. HarperCollins. p. 17. 6176:10.1016/S0169-5347(02)02490-4 5781:10.1016/s0047-2484(84)80079-2 5359:Reno PL, et al. (2010). 5051:Hooper, Rowan (31 May 2007). 4961:Hunt, Kevin (February 1996). 4696:10.1016/s0047-2484(84)80079-2 2637:The Secret World of Kangaroos 2559: 2265: 1973:A related study conducted by 818:Evolution of human bipedalism 695:Limited bipedalism in mammals 378:The first known biped is the 235: 216:The word is derived from the 6795:Human Timeline (Interactive) 6214:Trevathan, Wenda R. (1996). 5890:10.1016/j.jhevol.2016.02.006 5851:10.1016/0047-2484(91)90003-e 5690:Proc. Natl. Acad. Sci. U.S.A 5614:(2nd ed.). p. 235. 5589:10.1126/science.211.4480.341 5451:Reno PL, Lovejoy CO (2015). 5430:10.1016/j.jhevol.2009.01.002 5030:Kaplan, Matt (31 May 2007). 4988:Gibbons, Ann (31 May 2007). 4937:10.1016/j.jhevol.2013.10.002 4714:Shreeve, James (July 1996). 4590:10.1590/S1516-89132015060399 4514:10.1590/S1516-89132015060399 4446:10.1590/S1516-89132015060399 3422:10.1016/j.jhevol.2005.12.011 3084:10.1126/science.290.5493.969 3038:. 2000-11-03. Archived from 2585:National Wildlife Federation 274:endurance running hypothesis 211: 7: 8019:Evolutionary biology Portal 6957:Comparative foot morphology 6660:", Murray (London), (1871). 6556:. New York: Harper Collins. 6386:McMahon, Thomas A. (1984). 6086:Cunnane, Stephen C (2005). 3754:Waldman, Dan (2004-07-21). 3147:Penn State (1 March 2005). 2513: 2278: 1946:Postural feeding hypothesis 1752: 974: 595: 305: 249:), combat (in bears, great 228:'foot', as contrasted with 101:, meaning 'two feet' (from 10: 8101: 6761:Journal of Human Evolution 6709:Journal of Human Evolution 6695:Journal of Human Evolution 6291:10.1016/j.jtbi.2006.04.016 6004:The Aquatic Ape Hypothesis 5009:Minkel, JR (31 May 2007). 4917:Journal of Human Evolution 4855:Journal of Human Evolution 4676:Journal of Human Evolution 4189:10.1016/j.jtbi.2006.04.016 3593:10.1038/s41586-022-04901-z 3402:Journal of Human Evolution 3134:10.1016/j.crpv.2005.09.002 3105:Hutchinson, J. R. (2006). 2770:Journal of Zoology, London 2573:Stewart, D. (2006-08-01). 2464: 2373:A group of children racing 2364: 2299: 2042:Australopithecus afarensis 2036:Australopithecus afarensis 1957:Australopithecus afarensis 1930:Australopithecus africanus 1920:Australopithecus afarensis 1869:Australopithecus afarensis 1865:Australopithecus anamensis 1851: 1799:Australopithecus africanus 1765: 528: 507:. Some extinct members of 374:Early reptiles and lizards 18: 7984: 7964:Evolutionary anthropology 7941: 7915: 7868: 7806: 7725: 7684: 7677: 7623: 7547: 7470: 7395: 7349: 7342: 7325: 7285: 7221: 7192: 7156: 7147: 7114: 7099: 7036: 7010: 6947: 6902: 6856: 6846: 6840:Animal locomotion on land 5808:10.1017/s0140525x00079218 5072:Thorpe, Susannah (2007). 4348:10.1007/s00114-009-0637-3 4322:Niemitz, Carsten (2010). 2005:Sahelanthropus tchadensis 1898:turnover pulse hypothesis 6790:The Origin of Bipedalism 6567:Bramble, Dennis (1983). 6552:DeSilva, Jeremy (2021). 6494:Pontzer, Herman (2012). 6189:DeSilva, Jeremy (2021). 6033:. Alpha Books. pp. 5082:University of Birmingham 4149:National Geographic News 3825:"Walking tall after all" 2535: 2133:Other behavioural models 1975:University of Birmingham 867: 399: 65:that usually moves in a 7870:Origin of modern humans 6741:American Anthropologist 6688:American Anthropologist 6670:, 145, 195–199, (1925). 6593:10.1126/science.6849136 5711:10.1073/pnas.0901280106 5538:10.1126/science.1175834 5330:10.1126/science.1175802 5272:10.1126/science.1175824 4824:10.1126/science.1140799 4759:10.1126/science.1227123 4716:"Sunset on the savanna" 4393:10.1002/ajpa.1330340105 4277:10.1126/science.1140799 4216:Anthropological Science 4210:Kimura, Tasuku (2019). 4016:McHenry, H. M. (2009). 3944:10.1126/science.1109616 3803:10.1126/science.1146580 3643:Smithsonian Institution 3461:10.1002/ajpa.1330440207 3299:10.1080/106351598260824 1739:P a r a n t h r o p u s 1598:Dispersal beyond Africa 139:(a group that includes 8075:Terrestrial locomotion 6920:Rectilinear locomotion 6729:10.1006/jhev.1993.1021 6697:, 26, 183–202, (1994). 6690:, 63, 687–710, (1961). 6428:10.1098/rsbl.2010.0294 6330:10.1098/rstb.2010.0112 6121:Am. J. Phys. Anthropol 5377:10.1098/rstb.2010.0086 5058:New Scientist Magazine 4894:. New York: Scribner. 4875:10.1006/jhev.1996.0034 3114:Comptes Rendus Palevol 2809:Sharp, N.C.C. (1997). 2719:10.1098/rsif.2018.0276 2575:"A Bird Like No Other" 2483: 2438: 2374: 2309: 2230: 2214:aquatic ape hypothesis 2210:aquatic ape hypothesis 2142:Thermoregulatory model 2099:Neanderthal extinction 870:−10 — 757:walk on its front legs 608: 566:kangaroo rats and mice 169:kangaroo rats and mice 135:period some groups of 47:terrestrial locomotion 39: 7878:Recent African origin 7116:Last common ancestors 6915:Undulatory locomotion 6626:Ruina.tam.cornell.edu 5612:Understanding Emotion 3829:Research Intelligence 3536:Kondō, Shirō (1985). 2478: 2436: 2372: 2341:whole-body kinetics. 2307: 1848:Savannah-based theory 960:−1 — 950:−2 — 940:−3 — 930:−4 — 920:−5 — 910:−6 — 900:−7 — 890:−8 — 880:−9 — 603: 503:lineages such as the 69:manner is known as a 29: 8047:Evolutionary biology 7893:Behavioral modernity 7883:Multiregional origin 7663:archaic Homo sapiens 7658:Homo heidelbergensis 7603:Red Deer Cave people 6500:Current Anthropology 5243:Ardipithecus ramidus 2966:Journal of Mammalogy 2455:verbal communication 2093:Ardipithecus ramidus 2066:Ardipithecus ramidus 2056:Ardipithecus ramidus 2011:Ardipithecus ramidus 1554:Earliest stone tools 556:, most of which are 7530:H. neanderthalensis 7450:H. e. tautavelensis 6910:Concertina movement 6864:Arboreal locomotion 6721:1993JHumE..24..259J 6678:The Ancestor's Tale 6585:1983Sci...219..251B 6324:(1556): 3289–3299. 6283:2006JThBi.242..581S 5882:2016JHumE..94...72D 5843:1991JHumE..21..117W 5773:1984JHumE..13...91W 5702:2009PNAS..10614241K 5644:10.7554/eLife.44433 5581:1981Sci...211..341L 5530:2009Sci...326...74L 5509:Lovejoy CO (2009). 5422:2009JHumE..56..551H 5371:(1556): 3355–3363. 5322:2009Sci...326...75W 5264:2009Sci...326...94S 5210:10.1038/nature00879 5202:2002Natur.418..145B 5016:Scientific American 4929:2014JHumE..66...64P 4867:1996JHumE..30..389I 4816:2007Sci...316.1328T 4751:2012Sci...338..514G 4688:1984JHumE..13...91W 4625:2000Natur.404..382R 4582:2015arXiv150802739K 4506:2015arXiv150802739K 4467:Napier, JR (1964). 4438:2015arXiv150802739K 4340:2010NW.....97..241N 4328:Naturwissenschaften 4269:2007Sci...316.1328T 4263:(5829): 1328–1331. 4181:2006JThBi.242..581S 4114:10.1038/nature06342 4106:2007Natur.450.1075W 4100:(7172): 1075–1078. 3987:1988SciAm.259e.118L 3975:Scientific American 3795:2007Sci...318.1066C 3646:. August 14, 2016. 3585:2022Natur.609...94D 3487:"Coquerel's Sifaka" 3443:Rose, M.D. (1976). 3414:2006JHumE..50..552V 3255:National Geographic 3219:1993Natur.361...64S 3126:2006CRPal...5..519H 3076:2000Sci...290..969B 2924:1993FuEco...7..195D 2873:10.1038/nature03052 2865:2004Natur.432..345B 2675:10.1242/jeb.97.1.41 2248:obstetrical dilemma 2072:The Ancestor's Tale 1854:Savannah hypothesis 808:terrestrial animals 667:Injured individuals 660:Danuvius guggenmosi 539:giant ground sloths 7717:Self-domestication 7508:H. heidelbergensis 7457:H. e. yuanmouensis 7422:H. e. lantianensis 7149:Australopithecines 6133:10.1002/ajpa.21122 6027:Meier, R. (2003). 6006:. Souvenir Press. 5121:10.1002/ajpa.20284 4229:10.1537/ase.190219 3733:10.1007/BF02382940 3687:10.1002/ajpa.10058 3286:Systematic Biology 2912:Functional Ecology 2819:Journal of Zoology 2634:Penny, M. (2002). 2525:Orthograde posture 2484: 2439: 2375: 2310: 2291:Shoulder stability 1985:Provisioning model 1952:Indiana University 1355:H. heidelbergensis 745:giant ground sloth 690:Limited bipedalism 609: 606:Eadweard Muybridge 604:A Man Running; by 584:. One black bear, 458:Radiometric dating 189:australopithecines 40: 8028: 8027: 7969:Paleoanthropology 7911: 7910: 7888:Archaic admixture 7766:Stoned ape theory 7702:Endurance running 7619: 7618: 7615: 7614: 7611: 7610: 7466: 7465: 7429:H. e. nankinensis 7385:H. tsaichangensis 7321: 7320: 7059: 7058: 6943: 6942: 6579:(4582): 251–256. 6506:(S6): S346–S358. 6437:978-0-19-850022-3 6399:978-0-691-02376-2 6101:978-981-256-191-6 6072:978-3-406-51606-1 6044:978-0-02-864421-9 6013:978-0-285-63518-0 5988:on 26 March 2009. 5962:Hardy, Alister C. 5924:On Becoming Human 5575:(4480): 341–350. 5474:10.7717/peerj.925 5196:(6894): 145–151. 5167:978-0-07-340520-9 4901:978-0-684-80000-4 4810:(5829): 1328–31. 4745:(6106): 514–517. 4619:(6776): 382–385. 4035:978-0-674-03175-3 4018:"Human Evolution" 3547:978-4-13-066093-8 3070:(5493): 969–972. 2859:(7015): 345–352. 2713:(146): 20180276. 2647:978-0-7398-4986-6 2620:978-0-7876-5784-0 2580:National Wildlife 2482:- a bipedal robot 2334:center of gravity 2208:, as part of the 2169:natural selection 2031:sexual dimorphism 1820:sexual dimorphism 1790:sexual dimorphism 1764: 1763: 1756:million years ago 1715: 1714: 1693: 1692: 1671: 1670: 1663:Earliest rock art 1649: 1648: 1625: 1624: 1618:Earliest language 1605: 1604: 1585: 1584: 1561: 1560: 1541: 1540: 1531:Earliest sign of 1518: 1517: 1508:Earliest sign of 1495: 1494: 1475: 1474: 1455: 1454: 1435: 1434: 1078:Ou. macedoniensis 518:Effigia okeeffeae 296:Uner Tan syndrome 257:) or camouflage. 21:Bipedality (film) 8092: 8057: 8056: 8045: 8044: 8036: 8016: 8004: 8003: 7992: 7991: 7928:Human prehistory 7903:Recent evolution 7898:Early migrations 7840:Thermoregulation 7741:Expensive tissue 7712:Sexual selection 7682: 7681: 7554: 7553: 7436:H. e. pekinensis 7347: 7346: 7340: 7339: 7255:A. bahrelghazali 7224:Australopithecus 7154: 7153: 7124:Chimpanzee–human 7112: 7111: 7086: 7079: 7072: 7063: 7062: 6854: 6853: 6833: 6826: 6819: 6810: 6809: 6779: 6756: 6732: 6641: 6640: 6638: 6637: 6618: 6612: 6611: 6609: 6607: 6564: 6558: 6557: 6549: 6540: 6539: 6491: 6480: 6479: 6473: 6469: 6467: 6459: 6449: 6413: 6404: 6403: 6383: 6368: 6367: 6357: 6309: 6303: 6302: 6266: 6260: 6259: 6211: 6205: 6204: 6186: 6180: 6179: 6159: 6153: 6152: 6112: 6106: 6105: 6083: 6077: 6076: 6059:Niemitz, Carsten 6055: 6049: 6048: 6024: 6018: 6017: 5996: 5990: 5989: 5987: 5970: 5958: 5952: 5951: 5939: 5933: 5932: 5918: 5912: 5911: 5901: 5861: 5855: 5854: 5826: 5820: 5819: 5796:Behav. Brain Sci 5791: 5785: 5784: 5756: 5750: 5740: 5734: 5733: 5723: 5713: 5681: 5675: 5674: 5664: 5646: 5622: 5616: 5615: 5607: 5601: 5600: 5564: 5558: 5557: 5524:(5949): 74e1–8. 5515: 5506: 5497: 5496: 5486: 5476: 5455:Australopithecus 5448: 5442: 5441: 5405: 5399: 5398: 5388: 5356: 5350: 5349: 5305: 5299: 5298: 5296: 5249: 5236: 5230: 5229: 5187: 5178: 5172: 5171: 5147: 5141: 5140: 5100: 5094: 5093: 5091: 5078: 5069: 5063: 5062: 5048: 5042: 5041: 5027: 5021: 5020: 5006: 5000: 4999: 4995:Science Magazine 4985: 4979: 4978: 4958: 4949: 4948: 4912: 4906: 4905: 4885: 4879: 4878: 4850: 4844: 4843: 4799: 4793: 4792: 4786: 4778: 4734: 4728: 4727: 4722:. Archived from 4711: 4700: 4699: 4671: 4665: 4662: 4653: 4652: 4633:10.1038/35006045 4608: 4602: 4601: 4575: 4555: 4549: 4548: 4543:Day, MH (1986). 4540: 4534: 4533: 4499: 4479: 4473: 4472: 4464: 4458: 4457: 4431: 4411: 4405: 4404: 4376: 4370: 4369: 4359: 4319: 4313: 4312: 4248: 4242: 4241: 4231: 4207: 4201: 4200: 4164: 4158: 4156: 4151:. Archived from 4140: 4134: 4133: 4085: 4080:. Archived from 4069: 4063: 4062: 4046: 4040: 4039: 4013: 4007: 4006: 3970: 3964: 3963: 3927: 3921: 3920: 3918: 3917: 3902: 3896: 3895: 3893: 3892: 3886: 3873: 3867: 3866: 3861:. Archived from 3859:Tetrapod Zoology 3850: 3844: 3843: 3841: 3840: 3821: 3815: 3814: 3778: 3772: 3771: 3769: 3768: 3751: 3745: 3744: 3716: 3710: 3709: 3707: 3706: 3697:. Archived from 3666: 3660: 3659: 3657: 3655: 3634: 3628: 3627: 3625: 3579:(7925): 94–100. 3570: 3561: 3552: 3551: 3533: 3527: 3526: 3524: 3523: 3508: 3502: 3501: 3499: 3498: 3483: 3477: 3476: 3471:. Archived from 3440: 3434: 3433: 3393: 3387: 3386: 3384: 3383: 3368: 3362: 3361: 3359: 3358: 3318: 3312: 3311: 3301: 3277: 3271: 3270: 3268: 3267: 3258:. Archived from 3245: 3239: 3238: 3227:10.1038/361064a0 3202: 3196: 3189: 3183: 3182: 3167: 3161: 3160: 3144: 3138: 3137: 3120:(3–4): 519–530. 3111: 3102: 3096: 3095: 3057: 3051: 3050: 3048: 3047: 3028: 3022: 3021: 3019: 3012: 3003: 2997: 2996: 2994: 2963: 2950: 2944: 2943: 2907: 2901: 2900: 2850: 2841: 2835: 2834: 2813:Acinonyx jubatus 2806: 2800: 2799: 2797: 2796: 2790: 2784:. Archived from 2767: 2755: 2749: 2748: 2738: 2698: 2687: 2686: 2658: 2652: 2651: 2631: 2625: 2624: 2606: 2600: 2599: 2597: 2596: 2587:. Archived from 2570: 2553: 2546: 2492:robot locomotion 2380:thermoregulation 2330:spinal curvature 2225: 2198:marine biologist 2173:ecological niche 2077:Australopithecus 2020:Australopithecus 1963:Australopithecus 1834:Multiple factors 1780:Australopithecus 1744: 1742: 1741: 1727: 1725: 1709: 1700: 1695: 1687: 1685:Earliest clothes 1678: 1673: 1665: 1656: 1651: 1632: 1627: 1612: 1607: 1592: 1587: 1574:Earliest sign of 1568: 1563: 1548: 1543: 1533:Australopithecus 1525: 1520: 1502: 1497: 1488:Earliest bipedal 1482: 1477: 1468:Chimpanzee split 1462: 1457: 1442: 1437: 1422: 1417: 1403: 1402: 1388: 1387: 1371: 1357: 1343: 1315: 1302: 1282: 1269: 1241: 1239:Australopithecus 1228: 1213: 1196: 1183: 1159: 1146: 1133: 1120: 1107: 1094: 1082: 1063: 1050: 1037: 1025: 1011: 998: 985: 983: 971: 966: 961: 956: 951: 946: 941: 936: 931: 926: 921: 916: 911: 906: 901: 896: 891: 886: 881: 876: 871: 859: 852: 845: 839: 829: 828:Hominin timeline 822: 775:. Among the non- 721:Ground squirrels 678:and two chimps, 535:ground pangolins 423:Other archosaurs 360:Many species of 276:. Bipedality in 100: 99: 96: 95: 92: 89: 86: 83: 80: 8100: 8099: 8095: 8094: 8093: 8091: 8090: 8089: 8065: 8064: 8063: 8051: 8039: 8031: 8029: 8024: 7980: 7937: 7923:Human evolution 7907: 7864: 7808: 7802: 7781:Cooperative eye 7726:Specific models 7721: 7673: 7652:Homo antecessor 7607: 7543: 7537:H. rhodesiensis 7501:H. floresiensis 7462: 7443:H. e. soloensis 7415:H. e. georgicus 7391: 7355:H. gautengensis 7330: 7328: 7317: 7281: 7217: 7188: 7143: 7134:Orangutan–human 7103: 7095: 7093:Human evolution 7090: 7060: 7055: 7046:Fish locomotion 7032: 7006: 6939: 6898: 6884:Knuckle-walking 6842: 6837: 6786: 6772:Natural History 6650: 6648:Further reading 6645: 6644: 6635: 6633: 6620: 6619: 6615: 6605: 6603: 6565: 6561: 6550: 6543: 6492: 6483: 6471: 6470: 6461: 6460: 6438: 6414: 6407: 6400: 6384: 6371: 6310: 6306: 6267: 6263: 6212: 6208: 6201: 6187: 6183: 6160: 6156: 6113: 6109: 6102: 6084: 6080: 6073: 6056: 6052: 6045: 6025: 6021: 6014: 5997: 5993: 5985: 5968: 5959: 5955: 5950:(3–4): 213–236. 5944:Human Evolution 5940: 5936: 5919: 5915: 5862: 5858: 5827: 5823: 5792: 5788: 5757: 5753: 5743:Joseph Jordania 5741: 5737: 5696:(34): 14241–6. 5682: 5678: 5623: 5619: 5608: 5604: 5565: 5561: 5513: 5507: 5500: 5449: 5445: 5406: 5402: 5357: 5353: 5316:(5949): 75–86. 5306: 5302: 5294: 5258:(5949): 94–99. 5247: 5237: 5233: 5185: 5179: 5175: 5168: 5148: 5144: 5101: 5097: 5089: 5076: 5070: 5066: 5049: 5045: 5037:Nature Magazine 5028: 5024: 5007: 5003: 4986: 4982: 4959: 4952: 4913: 4909: 4902: 4886: 4882: 4851: 4847: 4800: 4796: 4780: 4779: 4735: 4731: 4712: 4703: 4672: 4668: 4663: 4656: 4609: 4605: 4556: 4552: 4541: 4537: 4480: 4476: 4465: 4461: 4412: 4408: 4377: 4373: 4320: 4316: 4249: 4245: 4208: 4204: 4165: 4161: 4141: 4137: 4078:The Independent 4070: 4066: 4055:Smithsonian.com 4047: 4043: 4036: 4014: 4010: 3971: 3967: 3928: 3924: 3915: 3913: 3904: 3903: 3899: 3890: 3888: 3884: 3874: 3870: 3865:on May 8, 2012. 3851: 3847: 3838: 3836: 3823: 3822: 3818: 3779: 3775: 3766: 3764: 3752: 3748: 3717: 3713: 3704: 3702: 3667: 3663: 3653: 3651: 3636: 3635: 3631: 3623: 3568: 3562: 3555: 3548: 3534: 3530: 3521: 3519: 3510: 3509: 3505: 3496: 3494: 3485: 3484: 3480: 3441: 3437: 3394: 3390: 3381: 3379: 3370: 3369: 3365: 3356: 3354: 3319: 3315: 3278: 3274: 3265: 3263: 3246: 3242: 3213:(6407): 64–66. 3203: 3199: 3190: 3186: 3169: 3168: 3164: 3145: 3141: 3109: 3103: 3099: 3058: 3054: 3045: 3043: 3030: 3029: 3025: 3017: 3010: 3004: 3000: 2992: 2978:10.2307/1381631 2961: 2955:Garland, T. Jr. 2951: 2947: 2932:10.2307/2389887 2908: 2904: 2848: 2842: 2838: 2807: 2803: 2794: 2792: 2788: 2765: 2759:Garland, T. Jr. 2756: 2752: 2699: 2690: 2659: 2655: 2648: 2632: 2628: 2621: 2607: 2603: 2594: 2592: 2571: 2567: 2562: 2557: 2556: 2547: 2543: 2538: 2516: 2473: 2465:Main articles: 2463: 2431: 2423:gluteus maximus 2404: 2396:Achilles tendon 2367: 2302: 2293: 2281: 2276: 2268: 2243: 2221: 2218:Carsten Niemitz 2190: 2181: 2179:Carrying models 2163:of early-stage 2144: 2135: 2118:warning display 2110:Joseph Jordania 2107: 2051: 2024:sexual swelling 1991:C. Owen Lovejoy 1987: 1948: 1939: 1856: 1850: 1836: 1812:knuckle-walking 1774: 1766:Main articles: 1760: 1759: 1747: 1746: 1745: 1737: 1736: 1734: 1730: 1729: 1728: 1723:H o m i n i d s 1721: 1719: 1711: 1710: 1705: 1698: 1689: 1688: 1683: 1676: 1667: 1666: 1661: 1654: 1645: 1644: 1630: 1621: 1620: 1610: 1601: 1600: 1590: 1581: 1580: 1575: 1566: 1557: 1556: 1546: 1537: 1536: 1523: 1514: 1513: 1500: 1491: 1490: 1480: 1471: 1470: 1460: 1451: 1450: 1440: 1431: 1430: 1420: 1413: 1412: 1411: 1406: 1405: 1404: 1398: 1397: 1393: 1391: 1390: 1389: 1383: 1382: 1378: 1376: 1375: 1374: 1367: 1361: 1360: 1359: 1353: 1348: 1346: 1345: 1344: 1335: 1327: 1319: 1318: 1311: 1305: 1304: 1303: 1294: 1286: 1285: 1278: 1272: 1271: 1270: 1261: 1253: 1245: 1244: 1237: 1231: 1230: 1229: 1220: 1216: 1215: 1214: 1205: 1201: 1200: 1199: 1192: 1186: 1185: 1184: 1175: 1167: 1163: 1162: 1161: 1155: 1150: 1149: 1148: 1142: 1137: 1136: 1135: 1129: 1124: 1123: 1122: 1116: 1111: 1110: 1109: 1103: 1098: 1097: 1096: 1092:Chororapithecus 1090: 1085: 1084: 1083: 1074: 1066: 1065: 1059: 1054: 1053: 1052: 1048:Samburupithecus 1046: 1041: 1040: 1039: 1033: 1028: 1027: 1026: 1019: 1015: 1014: 1013: 1007: 1002: 1001: 1000: 994: 989: 988: 987: 981: 979: 972: 969: 967: 964: 962: 959: 957: 954: 952: 949: 947: 944: 942: 939: 937: 934: 932: 929: 927: 924: 922: 919: 917: 914: 912: 909: 907: 904: 902: 899: 897: 894: 892: 889: 887: 884: 882: 879: 877: 874: 872: 869: 863: 837: 831: 827: 820: 796:basilisk lizard 792:monitor lizards 769: 697: 692: 684:knuckle-walking 598: 578:ground pangolin 531: 494:Middle Triassic 475:common ancestor 425: 402: 397: 376: 358: 356:Extant reptiles 342: 340:Bipedal animals 308: 286: 238: 214: 77: 73: 24: 17: 12: 11: 5: 8098: 8088: 8087: 8082: 8080:Animal anatomy 8077: 8062: 8061: 8049: 8026: 8025: 8023: 8022: 8010: 7998: 7985: 7982: 7981: 7979: 7978: 7973: 7972: 7971: 7961: 7956: 7951: 7945: 7943: 7939: 7938: 7936: 7935: 7933:Human timeline 7930: 7925: 7919: 7917: 7913: 7912: 7909: 7908: 7906: 7905: 7900: 7895: 7890: 7885: 7880: 7874: 7872: 7866: 7865: 7863: 7862: 7857: 7852: 7847: 7842: 7837: 7832: 7827: 7822: 7817: 7811: 7809: 7804: 7803: 7801: 7800: 7799: 7798: 7793: 7785: 7784: 7783: 7778: 7770: 7769: 7768: 7763: 7758: 7756:Drunken monkey 7750: 7749: 7748: 7743: 7738: 7729: 7727: 7723: 7722: 7720: 7719: 7714: 7709: 7704: 7699: 7694: 7688: 7686: 7685:General models 7679: 7675: 7674: 7672: 7671: 7629: 7627: 7621: 7620: 7617: 7616: 7613: 7612: 7609: 7608: 7606: 7605: 7600: 7595: 7590: 7585: 7578: 7573: 7564: 7562: 7551: 7545: 7544: 7542: 7541: 7533: 7526: 7519: 7511: 7504: 7497: 7489: 7484: 7476: 7474: 7472:Archaic humans 7468: 7467: 7464: 7463: 7461: 7460: 7453: 7446: 7439: 7432: 7425: 7418: 7411: 7403: 7401: 7393: 7392: 7390: 7389: 7381: 7377:H. rudolfensis 7373: 7366: 7359: 7350: 7344: 7337: 7323: 7322: 7319: 7318: 7316: 7315: 7308: 7301: 7298:P. aethiopicus 7293: 7291: 7283: 7282: 7280: 7279: 7272: 7265: 7258: 7251: 7244: 7237: 7229: 7227: 7219: 7218: 7216: 7215: 7208: 7200: 7198: 7190: 7189: 7187: 7186: 7179: 7176:Sahelanthropus 7172: 7165: 7162:Nakalipithecus 7157: 7151: 7145: 7144: 7142: 7141: 7136: 7131: 7126: 7120: 7118: 7109: 7097: 7096: 7089: 7088: 7081: 7074: 7066: 7057: 7056: 7054: 7053: 7051:Volant animals 7048: 7043: 7037: 7034: 7033: 7031: 7030: 7025: 7020: 7014: 7012: 7008: 7007: 7005: 7004: 6999: 6994: 6984: 6979: 6974: 6969: 6964: 6959: 6953: 6951: 6945: 6944: 6941: 6940: 6938: 6937: 6932: 6927: 6922: 6917: 6912: 6906: 6904: 6900: 6899: 6897: 6896: 6891: 6886: 6881: 6876: 6871: 6860: 6858: 6851: 6844: 6843: 6836: 6835: 6828: 6821: 6813: 6807: 6806: 6792: 6785: 6784:External links 6782: 6781: 6780: 6764: 6757: 6736: 6733: 6715:(4): 259–280. 6704: 6698: 6691: 6684: 6681: 6671: 6661: 6649: 6646: 6643: 6642: 6613: 6559: 6541: 6528:10.1086/667402 6512:10.1086/667402 6481: 6472:|journal= 6436: 6405: 6398: 6369: 6304: 6277:(3): 581–590. 6261: 6226:(2): 287–290. 6206: 6200:978-0062938497 6199: 6181: 6170:(5): 212–217. 6154: 6107: 6100: 6078: 6071: 6050: 6043: 6019: 6012: 6000:Morgan, Elaine 5991: 5953: 5934: 5913: 5856: 5837:(2): 117–136. 5821: 5786: 5751: 5735: 5676: 5617: 5602: 5559: 5498: 5443: 5416:(6): 551–559. 5400: 5351: 5300: 5231: 5173: 5166: 5142: 5115:(2): 225–231. 5095: 5064: 5043: 5022: 5001: 4980: 4950: 4907: 4900: 4880: 4861:(5): 389–397. 4845: 4794: 4729: 4726:on 2017-09-28. 4701: 4666: 4654: 4603: 4566:(6): 929–934. 4550: 4535: 4490:(6): 929–934. 4474: 4459: 4422:(6): 929–934. 4406: 4371: 4334:(3): 241–263. 4314: 4243: 4202: 4175:(3): 581–590. 4159: 4155:on 2008-09-11. 4135: 4084:on 2007-12-15. 4064: 4041: 4034: 4008: 3965: 3938:(5717): 1927. 3922: 3897: 3868: 3845: 3816: 3789:(5853): 1066. 3773: 3746: 3727:(4): 913–921. 3711: 3681:(2): 184–190. 3661: 3629: 3553: 3546: 3528: 3503: 3478: 3475:on 2013-01-05. 3455:(2): 247–261. 3435: 3408:(5): 552–567. 3388: 3363: 3333:(2): 463–478. 3313: 3292:(3): 457–474. 3272: 3240: 3197: 3193:Dinosaur Cards 3184: 3162: 3139: 3097: 3052: 3023: 2998: 2972:(4): 765–772. 2945: 2918:(2): 195–202. 2902: 2836: 2825:(3): 493–494. 2801: 2776:(2): 157–170. 2750: 2688: 2653: 2646: 2626: 2619: 2601: 2564: 2563: 2561: 2558: 2555: 2554: 2540: 2539: 2537: 2534: 2533: 2532: 2530:Quadrupedalism 2527: 2522: 2515: 2512: 2471:Chicken walker 2467:Humanoid robot 2462: 2461:Bipedal robots 2459: 2430: 2427: 2403: 2400: 2366: 2363: 2362: 2361: 2358: 2355: 2352: 2349: 2301: 2298: 2292: 2289: 2285:overcorrection 2280: 2277: 2275: 2272: 2267: 2264: 2242: 2239: 2189: 2186: 2180: 2177: 2159:environmental 2143: 2140: 2134: 2131: 2106: 2103: 2050: 2047: 1986: 1983: 1947: 1944: 1938: 1935: 1894:Elizabeth Vrba 1852:Main article: 1849: 1846: 1835: 1832: 1786:foramen magnum 1768:Human timeline 1762: 1761: 1753: 1749: 1748: 1733: 1732: 1731: 1718: 1717: 1716: 1713: 1712: 1704: 1703: 1701: 1691: 1690: 1682: 1681: 1679: 1669: 1668: 1660: 1659: 1657: 1647: 1646: 1636: 1635: 1633: 1623: 1622: 1616: 1615: 1613: 1603: 1602: 1596: 1595: 1593: 1583: 1582: 1572: 1571: 1569: 1559: 1558: 1552: 1551: 1549: 1539: 1538: 1529: 1528: 1526: 1516: 1515: 1506: 1505: 1503: 1493: 1492: 1486: 1485: 1483: 1473: 1472: 1466: 1465: 1463: 1453: 1452: 1446: 1445: 1443: 1433: 1432: 1426: 1425: 1423: 1414: 1409: 1408: 1407: 1396: 1395: 1394: 1392: 1381: 1380: 1379: 1377: 1364: 1363: 1362: 1351: 1350: 1349: 1347: 1308: 1307: 1306: 1290:H. rudolfensis 1275: 1274: 1273: 1234: 1233: 1232: 1219: 1218: 1217: 1204: 1203: 1202: 1189: 1188: 1187: 1166: 1165: 1164: 1153: 1152: 1151: 1144:Graecopithecus 1140: 1139: 1138: 1131:Sahelanthropus 1127: 1126: 1125: 1114: 1113: 1112: 1101: 1100: 1099: 1088: 1087: 1086: 1061:Ouranopithecus 1057: 1056: 1055: 1044: 1043: 1042: 1035:Nakalipithecus 1031: 1030: 1029: 1018: 1017: 1016: 1005: 1004: 1003: 992: 991: 990: 977: 976: 975: 973: 970:0 — 968: 963: 958: 953: 948: 943: 938: 933: 928: 923: 918: 913: 908: 903: 898: 893: 888: 883: 878: 873: 868: 865: 864: 862: 861: 854: 847: 836: 833: 832: 825: 819: 816: 768: 765: 737:quadrupedalism 696: 693: 691: 688: 669: 668: 654:Sahelanthropus 632:move like all 624:, gibbons and 597: 594: 537:, the extinct 530: 527: 473:resembles the 424: 421: 401: 398: 396: 393: 375: 372: 357: 354: 350:Tree kangaroos 341: 338: 337: 336: 330: 324: 318: 307: 304: 285: 282: 255:monitor lizard 253:and the large 247:giant pangolin 237: 234: 213: 210: 15: 9: 6: 4: 3: 2: 8097: 8086: 8083: 8081: 8078: 8076: 8073: 8072: 8070: 8060: 8055: 8050: 8048: 8043: 8038: 8037: 8034: 8021: 8020: 8015: 8011: 8009: 8008: 7999: 7997: 7996: 7987: 7986: 7983: 7977: 7974: 7970: 7967: 7966: 7965: 7962: 7960: 7957: 7955: 7952: 7950: 7947: 7946: 7944: 7940: 7934: 7931: 7929: 7926: 7924: 7921: 7920: 7918: 7914: 7904: 7901: 7899: 7896: 7894: 7891: 7889: 7886: 7884: 7881: 7879: 7876: 7875: 7873: 7871: 7867: 7861: 7858: 7856: 7853: 7851: 7848: 7846: 7843: 7841: 7838: 7836: 7833: 7831: 7828: 7826: 7823: 7821: 7818: 7816: 7813: 7812: 7810: 7805: 7797: 7794: 7792: 7789: 7788: 7787:Life history 7786: 7782: 7779: 7777: 7774: 7773: 7771: 7767: 7764: 7762: 7759: 7757: 7754: 7753: 7751: 7747: 7744: 7742: 7739: 7737: 7734: 7733: 7731: 7730: 7728: 7724: 7718: 7715: 7713: 7710: 7708: 7705: 7703: 7700: 7698: 7695: 7693: 7690: 7689: 7687: 7683: 7680: 7676: 7670: 7669: 7664: 7660: 7659: 7654: 7653: 7648: 7647: 7642: 7641: 7640:Homo ergaster 7636: 7635: 7631: 7630: 7628: 7626: 7622: 7604: 7601: 7599: 7596: 7594: 7591: 7589: 7586: 7584: 7583: 7579: 7577: 7574: 7572: 7570: 7569:H. s. sapiens 7566: 7565: 7563: 7561: 7560: 7555: 7552: 7550: 7549:Modern humans 7546: 7539: 7538: 7534: 7532: 7531: 7527: 7525: 7524: 7523:H. luzonensis 7520: 7517: 7516: 7512: 7510: 7509: 7505: 7503: 7502: 7498: 7495: 7494: 7490: 7488: 7485: 7483: 7482: 7481:H. antecessor 7478: 7477: 7475: 7473: 7469: 7459: 7458: 7454: 7452: 7451: 7447: 7445: 7444: 7440: 7438: 7437: 7433: 7431: 7430: 7426: 7424: 7423: 7419: 7417: 7416: 7412: 7410: 7409: 7408:H. e. erectus 7405: 7404: 7402: 7400: 7399: 7394: 7387: 7386: 7382: 7379: 7378: 7374: 7372: 7371: 7367: 7365: 7364: 7360: 7357: 7356: 7352: 7351: 7348: 7345: 7341: 7338: 7336: 7334: 7324: 7314: 7313: 7309: 7307: 7306: 7302: 7300: 7299: 7295: 7294: 7292: 7290: 7289: 7284: 7278: 7277: 7273: 7271: 7270: 7266: 7264: 7263: 7262:A. deyiremeda 7259: 7257: 7256: 7252: 7250: 7249: 7245: 7243: 7242: 7238: 7236: 7235: 7231: 7230: 7228: 7226: 7225: 7220: 7214: 7213: 7209: 7207: 7206: 7202: 7201: 7199: 7197: 7196: 7191: 7185: 7184: 7183:Kenyanthropus 7180: 7178: 7177: 7173: 7171: 7170: 7166: 7164: 7163: 7159: 7158: 7155: 7152: 7150: 7146: 7140: 7137: 7135: 7132: 7130: 7129:Gorilla–human 7127: 7125: 7122: 7121: 7119: 7117: 7113: 7110: 7107: 7102: 7098: 7094: 7087: 7082: 7080: 7075: 7073: 7068: 7067: 7064: 7052: 7049: 7047: 7044: 7042: 7039: 7038: 7035: 7029: 7026: 7024: 7021: 7019: 7016: 7015: 7013: 7009: 7003: 7000: 6998: 6995: 6992: 6988: 6985: 6983: 6980: 6978: 6975: 6973: 6970: 6968: 6965: 6963: 6962:Arthropod leg 6960: 6958: 6955: 6954: 6952: 6950: 6946: 6936: 6933: 6931: 6928: 6926: 6923: 6921: 6918: 6916: 6913: 6911: 6908: 6907: 6905: 6901: 6895: 6892: 6890: 6887: 6885: 6882: 6880: 6877: 6875: 6872: 6869: 6865: 6862: 6861: 6859: 6855: 6852: 6849: 6845: 6841: 6834: 6829: 6827: 6822: 6820: 6815: 6814: 6811: 6805:(August 2016) 6804: 6800: 6796: 6793: 6791: 6788: 6787: 6777: 6773: 6769: 6765: 6762: 6758: 6754: 6750: 6746: 6742: 6737: 6734: 6730: 6726: 6722: 6718: 6714: 6710: 6705: 6702: 6699: 6696: 6692: 6689: 6685: 6682: 6679: 6675: 6672: 6669: 6665: 6662: 6659: 6655: 6652: 6651: 6631: 6627: 6623: 6617: 6602: 6598: 6594: 6590: 6586: 6582: 6578: 6574: 6570: 6563: 6555: 6548: 6546: 6537: 6533: 6529: 6525: 6521: 6517: 6513: 6509: 6505: 6501: 6497: 6490: 6488: 6486: 6477: 6465: 6457: 6453: 6448: 6443: 6439: 6433: 6429: 6425: 6421: 6420: 6412: 6410: 6401: 6395: 6391: 6390: 6382: 6380: 6378: 6376: 6374: 6365: 6361: 6356: 6351: 6347: 6343: 6339: 6335: 6331: 6327: 6323: 6319: 6315: 6308: 6300: 6296: 6292: 6288: 6284: 6280: 6276: 6272: 6265: 6257: 6253: 6249: 6245: 6241: 6237: 6233: 6229: 6225: 6221: 6217: 6210: 6202: 6196: 6192: 6185: 6177: 6173: 6169: 6165: 6158: 6150: 6146: 6142: 6138: 6134: 6130: 6127:(4): 630–42. 6126: 6122: 6118: 6111: 6103: 6097: 6093: 6089: 6082: 6074: 6068: 6064: 6060: 6054: 6046: 6040: 6036: 6032: 6031: 6023: 6015: 6009: 6005: 6001: 5995: 5984: 5980: 5976: 5975: 5974:New Scientist 5967: 5963: 5957: 5949: 5945: 5938: 5930: 5926: 5925: 5917: 5909: 5905: 5900: 5895: 5891: 5887: 5883: 5879: 5875: 5871: 5867: 5860: 5852: 5848: 5844: 5840: 5836: 5832: 5825: 5817: 5813: 5809: 5805: 5801: 5797: 5790: 5782: 5778: 5774: 5770: 5766: 5762: 5755: 5749:. 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I. 6694: 6687: 6667: 6634:. Retrieved 6625: 6616: 6604:. Retrieved 6576: 6572: 6562: 6553: 6503: 6499: 6418: 6388: 6321: 6317: 6307: 6274: 6270: 6264: 6223: 6219: 6209: 6190: 6184: 6167: 6163: 6157: 6124: 6120: 6110: 6087: 6081: 6062: 6053: 6029: 6022: 6003: 5994: 5983:the original 5978: 5972: 5956: 5947: 5943: 5937: 5923: 5916: 5873: 5870:J. Hum. Evol 5869: 5859: 5834: 5831:J. Hum. Evol 5830: 5824: 5799: 5795: 5789: 5767:(1): 91–98. 5764: 5761:J. Hum. Evol 5760: 5754: 5738: 5693: 5689: 5679: 5634: 5630: 5620: 5611: 5605: 5572: 5568: 5562: 5521: 5517: 5464: 5460: 5454: 5446: 5413: 5409: 5403: 5368: 5364: 5354: 5313: 5309: 5303: 5255: 5251: 5242: 5234: 5193: 5189: 5176: 5152: 5145: 5112: 5108: 5098: 5080: 5067: 5056: 5046: 5035: 5025: 5014: 5004: 4993: 4983: 4970: 4966: 4920: 4916: 4910: 4890: 4883: 4858: 4854: 4848: 4807: 4803: 4797: 4783:cite journal 4742: 4738: 4732: 4724:the original 4719: 4682:(1): 91–98. 4679: 4675: 4669: 4616: 4612: 4606: 4563: 4559: 4553: 4544: 4538: 4487: 4483: 4477: 4468: 4462: 4419: 4415: 4409: 4387:(1): 55–60. 4384: 4380: 4374: 4331: 4327: 4317: 4260: 4256: 4246: 4219: 4215: 4205: 4172: 4168: 4162: 4153:the original 4148: 4138: 4097: 4093: 4082:the original 4077: 4067: 4054: 4044: 4021: 4011: 3981:(5): 82–89. 3978: 3974: 3968: 3935: 3931: 3925: 3914:. Retrieved 3900: 3889:. Retrieved 3882:the original 3871: 3863:the original 3858: 3848: 3837:. 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Retrieved 2589:the original 2578: 2568: 2550:red kangaroo 2544: 2485: 2451: 2440: 2408:domesticated 2405: 2392:plantar arch 2389: 2376: 2343: 2338:Force plates 2319: 2314: 2311: 2294: 2282: 2274:Biomechanics 2269: 2252: 2244: 2241:Consequences 2235: 2191: 2182: 2149:Ardipithecus 2145: 2136: 2122:intimidation 2108: 2091: 2083:Paranthropus 2081: 2075: 2064: 2054: 2052: 2041: 2034: 2028: 2019: 2016:Ardipithecus 2015: 2009: 2003: 1988: 1972: 1961: 1955: 1949: 1940: 1928: 1918: 1912: 1903: 1888: 1884: 1881:synapomorphy 1876: 1872: 1868: 1864: 1857: 1839: 1837: 1808: 1797: 1794:lumbar spine 1778: 1775: 1735: 1720: 1576: 1532: 1510:Ardipithecus 1509: 1428:Earlier apes 1385:Neanderthals 1369:Homo sapiens 1366: 1365: 1352: 1336: 1328: 1320: 1310: 1309: 1295: 1287: 1277: 1276: 1262: 1254: 1246: 1236: 1235: 1221: 1206: 1194:Ardipithecus 1191: 1190: 1176: 1168: 1154: 1141: 1128: 1118:Sivapithecus 1115: 1105:Oreopithecus 1102: 1089: 1075: 1067: 1058: 1045: 1032: 1020: 1006: 993: 978: 770: 698: 673: 670: 658: 652: 642: 610: 570:hopping mice 551: 532: 516: 509:Pseudosuchia 498: 484: 478: 470: 464: 443: 436: 433:crocodilians 426: 403: 382: 377: 365: 359: 343: 309: 299: 287: 266:red kangaroo 259: 239: 225: 221: 215: 191:, including 177:hopping mice 130: 122:bipedal gait 121: 109: 105: 70: 66: 42: 41: 36:red kangaroo 7791:Grandmother 7746:Shore-based 7707:Aquatic ape 7598:Tam Pa Ling 7493:H. ergaster 7312:P. robustus 7018:Canine gait 6991:Facultative 6977:Unguligrade 6972:Plantigrade 6967:Digitigrade 6935:Other modes 6930:Sidewinding 6868:Brachiation 6799:Smithsonian 6778:(5): 47–51. 6674:Dawkins, R. 6664:Dart, R. A. 4222:(1): 1–12. 2429:Respiration 2402:Musculature 2260:birth canal 2114:aposematism 1925:Little Foot 1331:H. ergaster 1224:Ar. ramidus 1209:Ar. kadabba 1171:O. praegens 1009:Pleistocene 838:This box: 804:cockroaches 761:anal glands 614:Chimpanzees 558:quadrupedal 505:iguanodonts 491:Argentinian 415:. However, 8085:2 (number) 8069:Categories 7830:Skin color 7815:Bipedalism 7776:Killer ape 7588:Cro-Magnon 7487:Denisovans 7363:H. habilis 7327:Humans and 7212:A. ramidus 7205:A. kadabba 7028:Human gait 7023:Horse gait 6747:(6): 738. 6654:Darwin, C. 6636:2013-04-30 5802:(2): 366. 5637:: e44433. 5410:J Hum Evol 5245:dentition" 4573:1508.02739 4497:1508.02739 4429:1508.02739 3916:2013-04-30 3891:2007-10-29 3839:2013-04-30 3767:2007-10-29 3705:2013-04-30 3654:August 14, 3522:2012-07-23 3497:2009-06-15 3382:2023-01-06 3357:2011-10-18 3266:2007-10-29 3046:2007-10-17 2795:2010-04-11 2595:2014-05-30 2560:References 2442:Quadrupeds 2415:quadriceps 2266:Physiology 1400:Denisovans 1339:Au. sediba 1313:H. erectus 1280:H. habilis 1070:Ou. turkae 800:arthropods 773:amphibians 649:bipedalism 590:Derbyshire 574:springhare 523:Pterosaurs 486:Lagerpeton 480:Marasuchus 429:archosaurs 380:bolosaurid 367:Ctenosaura 236:Advantages 224:'two' and 173:springhare 141:crocodiles 137:archosaurs 43:Bipedalism 7949:Theorists 7916:Timelines 7796:Patriarch 7772:Behavior 7697:Gathering 7625:Ancestors 7370:H. naledi 7305:P. boisei 7276:A. sediba 7002:Quadruped 6606:28 August 6520:0011-3204 6474:ignored ( 6464:cite book 6338:0962-8436 6240:0745-5194 5876:: 72–82. 5816:147314740 5653:2050-084X 5129:0002-9483 4973:: 77–90. 4923:: 64–82. 4775:206543814 4285:0036-8075 4238:132162687 3811:0036-8075 3617:234630242 3601:0028-0836 2881:1476-4687 2727:1742-5689 2669:: 41–56. 2520:Allometry 2419:hamstring 2385:endurance 2223:‹See Tfd› 2088:hominines 2061:homininae 1995:evolution 1979:orangutan 1860:hominines 1804:pregnancy 1298:Au. garhi 777:archosaur 562:macropods 547:macropods 446:archosaur 413:dinosaurs 384:Eudibamus 230:quadruped 212:Etymology 181:pangolins 165:macropods 159:. Within 157:theropods 149:dinosaurs 145:dinosaurs 7995:Category 7850:Language 7820:Skeleton 7515:H. longi 7269:A. garhi 7106:Hominins 7101:Taxonomy 7011:Specific 6768:Vrba, E. 6630:Archived 6536:31461168 6456:20410030 6364:20855303 6346:20778968 6299:16782133 6149:36325131 6141:19890871 6065:. Beck. 6061:(2004). 6002:(1997). 5964:(1960). 5929:Archived 5908:27178459 5730:19667206 5671:31038121 5597:17748254 5554:42790876 5546:19810200 5493:25945314 5467:. e925. 5438:19446306 5395:20855309 5346:20189444 5338:19810190 5292:Archived 5280:19810195 5218:12110880 5137:16288480 5087:Archived 4975:Archived 4945:24315239 4840:85992565 4832:17540902 4767:23112331 4720:Discover 4641:10746723 4366:20127307 4309:85992565 4301:17540902 4293:20036393 4197:16782133 4122:18075592 4086:Quoting 4059:Archived 3960:21030132 3952:15790846 3910:Archived 3761:NBC News 3741:41892278 3721:Primates 3695:12012370 3648:Archived 3621:Archived 3609:36002567 3516:Archived 3430:16516949 3376:BBC News 3351:Archived 3347:84129882 3308:12066687 3179:Archived 3157:Archived 3092:11062126 3015:Archived 2990:Archived 2957:(1988). 2889:15549097 2761:(1983). 2745:30257922 2514:See also 2394:and the 2279:Standing 2157:woodland 2090:such as 1999:hominids 1816:grasping 1022:Hominini 996:Pliocene 786:such as 780:reptiles 725:meerkats 705:raccoons 622:gorillas 596:Primates 513:Triassic 501:dinosaur 471:Eoraptor 466:Eoraptor 450:Triassic 438:Eoraptor 410:theropod 315:Standing 306:Movement 300:normally 264:and the 206:indriids 133:Triassic 8059:Science 8033:Portals 8007:Commons 7959:Fossils 7825:Muscles 7736:Cooking 7692:Hunting 7169:Orrorin 6949:Anatomy 6925:Rolling 6903:Legless 6894:Walking 6889:Running 6879:Jumping 6717:Bibcode 6601:6849136 6581:Bibcode 6573:Science 6447:2880073 6355:2981964 6279:Bibcode 6256:8744088 5899:4874949 5878:Bibcode 5839:Bibcode 5769:Bibcode 5721:2732797 5698:Bibcode 5662:6491036 5577:Bibcode 5569:Science 5526:Bibcode 5518:Science 5484:4419524 5418:Bibcode 5386:2981962 5318:Bibcode 5310:Science 5288:3744438 5260:Bibcode 5252:Science 5226:1316969 5198:Bibcode 4925:Bibcode 4863:Bibcode 4812:Bibcode 4804:Science 4747:Bibcode 4739:Science 4684:Bibcode 4649:4303978 4621:Bibcode 4578:Bibcode 4522:4993117 4502:Bibcode 4434:Bibcode 4401:4993117 4357:2819487 4336:Bibcode 4265:Bibcode 4257:Science 4177:Bibcode 4102:Bibcode 4003:3212438 3983:Bibcode 3932:Science 3791:Bibcode 3783:Science 3581:Bibcode 3410:Bibcode 3235:4270484 3215:Bibcode 3177:. 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