The Origins of Bipedalism - discuss various theories that attempt to explain how and

why humans began to walk on two legs.

Atika Farooq
0081868
30, November 2023
 Introduction

Bipedalism, the ability to walk on two legs made a significant change in human evolution. It is a

primary trait that sets hominins unique from other primates. These adaptions provide flexible

hand mobility for daily tasks like carrying and using tools (PBS Eons, 2018). It is thought that

this enormous evolutionary leap is what set off a variety of skeletal changes shared by all bipedal

hominins over the century. Various hypotheses have been proposed to demonstrate the evolution

of bipedalism in humans. One of them is the Savannah hypothesis, which provides an in-

depth discussion of why we became biped in the first place. Why did the development of

bipedalism occur in the first place? Other conventional theories also suggest that bipedalism

evolved from a knuckle-walking terrestrial phase or originated in arboreal environments as a

vertical climber (Thorpe et al., 2014:906-907) (Burke et al. 2019). This article will explore a

variety of theories that explain the Journey of bipedalism and energy Expenditure. starting with

Charles Darwin's discovery of bipedalism as a critical aspect of human evolution as he proposed

that the evolution of bipedalism allowed our ancestors to carry out tasks in addition to

locomotion. which evaluates fossil evidence, examines the anatomical transformation, and

discusses the evolutionary implications of this major leap further in the evolution of humans

(Rodman & McHenry,1980:103-105).

 Theories of Bipedalism

Traditionally, bipedalism has been seen as one of the defining traits of our lineage. The development

of longitudinal arches in the human foot is set to be the most notable phylogenetic difference

between humans and other primates in terms of foot evolution (PBS Eons, 2018). One of the

Energy Expenditure Theories suggests that the evolution of bipedalism, or walking on two legs,

was driven by a change in how food resources were spread out through the environment. This

change required a more energy-efficient way of moving around. Several studies have analyzed

the energy used by different types of movement in humans and apes, and these have been

supported by this theory. The theory also suggests that changes in the environment and

availability of resources had a huge impact on the evolution of bipedalism than changes in food

or social structures. specifically, during the late Miocene period dated back to between 5 to 20

Mys, resources became more scattered due to the emergence of open habitats. (Roberts &

Thorpe, 2014). To make the most of these resources, early human ancestors needed a more

efficient way of locomotion than the quadrupedal walking gait pattern of movement often visible

by apes like chimpanzees), this theory was supported by the study done by Rodman and

McHenry as they measured the oxygen and energy used by human and chimpanzees while

walking or running. According to their research, humans use significantly less energy while

walking at higher speeds compared to chimps. They estimated that if a human walks at a speed

of 5 kilometres per hour, they would need only one-fourth of the energy required by a

chimpanzee to travel at the same speed. They also estimated that humans could cover 80

kilometres in a day, using the same amount of energy a chimp would need to travel just 27
kilometres (Rodman & McHenry 1980:103-105). Another research study done by Pontzer and

his team supported this concept by Using a method known as doubly labelled water, they

calculated the energy expenditure of apes and humans in their natural habitats (PBS Eons, 2018).

They discovered that despite having comparable body proportions and levels of activity, humans

consume less energy daily than chimpanzees and orangutans. Moving around requires less

energy in humans than in apes. The primary reason for this variation is that two-legged walking

is more efficient (Burke et al. 2019). They concluded that bipedalism possibly had a significant

role in human evolution by reducing the energy requirements of long-distance travel. To

understand how we began to walk on two feet (Lupu et al. 2017:58-59). It is crucial to first

identify the development of human anatomy. The discovery of a 4.4-million-year-old

Ardipithecus ramidus skeleton in Ethiopia in 1994 transformed our understanding of the

transitional period, which was commonly known as Ardi, and has led scientists to believe that

hominids (a group that includes humans and their common ancestors with chimpanzees) did not

undergo an intermediate stage of knuckle-walking before transitioning to bipedalism (Rodman &

McHenry,1980:103-105). During the 1970s, paleoanthropologist Russell H. Tuttle (1974, 1981)

proposed that the arboreal ancestor of modern hominins would have been small around the size

of living gibbons weighing 9–13.5kg and would have engaged extensively in vertical climbing,

which is functionally associated with bipedalism. Suggesting from her unique features Ardi was

from a woodland environment indicating she might be both a climber and walker on the ground.

However, when considering Ardipithecus’ locomotion (PBS Eons, 2018). According

to Crompton et al (2010), Ardipithecus' huge body size and stiffness, and non-compliant feet

were not ideal for movements around the trees. however, there are still arguments over

Ardipithecus's possible tree movement, it is generally agreed that this species was terrestrially
bipedal on the ground. While there isn't any sufficient evidence supporting that Ardipithecus was

a knuckle-walker or that it descended from a previous knuckle-walker (Lupu et al., 2017:58-59).

Additionally, the footprint discovered at the Laetoli site in Tanzania back to 3.6 Mys made by

Australopithecus afarnesis another early human ancestor suggests that walking upright may have

been what initially set our lineage apart rather than intelligence. Now, the fact that

Australopithecus was able to walk upright around 3.6 million years ago raises questions

concerning the origins of this type of locomotion. Which species was the first to follow this

path? Who made the first moves toward what we now call walking? According to some

Paleoanthropologists, there were some early ancestors in line to be considered as the first to

walk. One of them was Australopithecus anamensis, which came from Africa between 4.2 and

3.9 million years ago. The second is Orrorin Tugenesnis around 6 million Mys found in the

Tugen Hills region of central Kenya (Senut et al., 2018:286). Lastly, the oldest biped

Sahelanthropus tchadensis which viewed as biped due to the hole position on the cranial base.

Interestingly, Ardipithecus kadabba an older species of Ardipithecus discovered in the Middle

Awash region of Ethiopia suggests it may be biped because the shape of the joint of bone of the

big toe was like that of a human (PBS Eons, 2018). However, due to little evidence of fossil

species being found, there have been debates regarding whether it has evidence of bipedalism. Or

which one was the first biped? Most agreed with Australopithecus anamensis as being the one

and others would argue with Ardi. The transformation from climbing trees to walking upright on

the ground played a vital step in human evolution (Thorpe et al., 2014:906-907) (Richmond,

Begun, and Strait, 2001).

The Savannah Hypothesis explains how early hominids evolved to walk on two feet as they

ventured out of the forest and into the savannah. As the environment changed, walking upright
became an advantageous adaptation that allowed for the acquisition of new skills that led to

better chances of survival (Senut et al., 2017:286). This evolutionary development provided a

unique advantage to early humans, enabling them to better navigate their surroundings and

explore new territories (Popwich, 2021). According to this hypothesis, the African savannah

climate played a huge impact as it led to the shrinking of forests and the growth of grassland,

often leading to a variety of theories and models like bipedalism the act of walking upright. As

the landscape changed. The upright posture also reduces the amount of body surface exposed to

the sun helping us regulate body temperature in savannah climate (Senut et al., 2017:286). This

also led to free movement of our hands allowing us to carry items and perform which was seen

as a crucial step in our evolution. A concept that inspired Darwin's theories. (Popwich, 2021)

Darwin (1871) added to this theory by arguing that our ancestors lived in trees and that the

knuckle-walking behaviour observed in chimpanzees and gorillas originated after humans

diverged from them. similarly, their evolutionary process would have changed if they had gone

from living in trees to living on the ground. This transition would let the animal become more

exclusively quadrupedal or bipedal because of this (Thorpe et al., 2014:906-907).

Evidence

Fossilized remains of early hominids, such as the Laetoli footprints Ardipithecus

ramidus (Ardi), and Australopithecus afarensis (Lucy), provide significant insight into the early

stages of bipedalism. These fossils provide important evidence that either confirms or disproves

the energy expenditure theory. They specifically highlight the physical distinctions between

bipedal hominins and their quadrupedal ape relatives about the pelvis, foot, and toes. These

distinctions highlight the special adaptations that allowed our ancestors to walk on two feet

(Rodman & McHenry, 1980:103-105). As I mentioned earlier, one of the earliest fossils that
demonstrate the evolution of bipedalism in early humans is that of Ardipithecus ramidus, which

was found in Africa in the late 20th century. These remains, dated back to 4.4 million years old,

are believed to represent a creature close to the last common ancestor between humans and

chimpanzees (Burke et al., 2019). Analysis of the fossil revealed that Ardipithecus had a big toe

that was designed for climbing, but it also had a pelvis and feet that were adapted for bipedal

locomotion. The pelvis of Ardipithecus is shorter and broader than that of apes as she was only

1.2 meters, which allows for a more upright posture and a more efficient transfer of weight from

the spine to the legs. The feet of Ardipithecus are also more rigid and arched providing more

stability and propulsion during walking. These features suggest that Ardipithecus was capable of

both climbing and walking on two legs, with this evidence we can tell that, Bipedalism was

already a characteristic of the early human lineage (Burke et al., 2019). In addition, the discovery

of a female Australopithecus afarensis fossil, known as Lucy, in Ethiopia in 1974 has also

provided further insights into early bipedalism. This shows a 3.2-million-year-old fossil exhibits

characteristics indicative of early bipedal development, close to certain ape-like features. Lucy’s

pelvis, while like that of Ardipithecus, is more forward-leaning, presenting a more consistent

bipedal stance. Her feet, more akin to those of humans than apes, feature a large toe aligned with

the other toes and a larger, more muscular heel. Despite these adaptations, Lucy retained a small

brain, a long, curved spine, and shoulders adapted for climbing. These traits suggest that while

Lucy spent a considerable amount of time on the ground, she was still primarily arboreal. This

combination of features underscores the complex evolutionary path toward bipedalism (Rodman

& McHenry, 1980:103-105). Another previously mentioned vitally important fossil find that

supports the energy expenditure and savannah is the theory of energy expenditure supported by

the discovery of the Laetoli footprints in Tanzania in 1978. They proposed that these footprints
were left by three human ancestors who walked through wet volcanic ash 3.6 million years ago.

The pattern of the footprints shows that the main stress was on the ball and heel of the foot, with

secondary stress on the outside of the foot in line with the toe. This way of walking is like

modern human's bipedal walking pattern and is unlike the flat-footed and inward-angled walking

of apes (PBS Eons, 2018). The fossil also reveals that the human ancestors had a fully developed

bipedal gait, with stride length and width like that of modern humans. The human ancestors

walked at a speed of approximately 1.5 m/s, indicating that they were able to travel long

distances efficiently. The discovery of Lucy, Ardipithecus, and the Laetoli footprints provide

evidence to support the theory that early humans evolved to walk on two feet because of changes

in their environment and the availability of resources. This required the development of a more

energy-efficient mode of transportation. These fossils also demonstrate that the evolution of

bipedalism was a gradual process and that it coexisted with other adaptations such as climbing

and brain growth (Roberts & Thorpe, 2014). However, the study of human evolution by Charles

Darwin on bipedalism theorized that our forebears were originally arboreal, and the knuckle-

walking seen in chimpanzees and gorillas evolved after our lineage split from theirs. He

speculated that if our ancestors had moved from an arboreal to a terrestrial lifestyle, the course of

evolution might have been different. (Rodman & McHenry, 1980:103-105) (Richmond et. al,

2001:73-75).

Modern research studied on Darwin’s theories, has reshaped our comprehension of the origins of

bipedalism. However, recent discoveries have questioned some of Darwin’s initial thoughts,

especially concerning the timeline of bipedal evolution. Bipedalism emerged at least 4.2 million

years ago, or possibly as far back as 6 million years ago, which is considerably earlier than the
estimated age of the earliest tools. This raises questions about the original purpose of bipedalism

(Thorpe et al. 2014:906-907) (Darwin 1871). As scientists investigated early hominid fossils

more thoroughly, the reasoning behind the evolution of bipedalism was questioned. While

Darwin’s theory had its strengths, the postural feeding hypothesis emerged as an alternative

explanation for the development of bipedalism. This theory posits that early hominins exhibited

partial bipedalism as they adapted to terrestrial life before becoming fully bipedal. The postural

feeding hypothesis, explored by researchers, suggests that the transition to bipedalism was more

closely linked to changes in feeding habits than to tool use. The theory argues that early

hominids adopted a more upright posture to access food resources more efficiently, which altered

their locomotion. Interestingly, the shift from Darwinian theory to the postural feeding

hypothesis shows the evolving nature of scientific inquiry and highlights the factors influencing

bipedalism. (Thorpe et al., 2014:906-907)

Conclusion

In conclusion, the development of bipedalism has been a crucial factor in shaping human

history and distinguishing us from our ape ancestors. Through various perspectives, including the

Savannah Hypothesis, Energy Expenditure theory, fossil records, anatomical modifications, and

theories ranging from Darwin's observations to the postural feeding hypothesis, we have gained a

multifaceted understanding of the process leading to bipedalism (Lupu et al. 2017:58-59.) The

anatomical adaptations and fossils like Australopithecus afarensis (Lucy) and Ardipithecus

ramidus (Ardi) demonstrate the progressive nature of bipedal development and its coexistence

with other features. The Laetoli footprints further emphasize how external factors have

influenced our ancestors' locomotive methods and solidify the switch to an energy-efficient
bipedal walk (Thorpe et al., 2014:906-907). The postural feeding hypothesis challenges the

traditional narrative and highlights the importance of eating habits in the adoption of an upright

posture (Wayman 2012). The dynamic nature of scientific investigation continually enhances our

understanding, delaying the birth of bipedalism and raising deeper questions about its initial

function (PBS Eons, 2018). Overall, bipedalism's evolutionary path is a complex tapestry of

adaptation that has improved mobility, responded to environmental changes, and contributed to

the unique course of human development. In addition, the lack of evidence, and discovery of

new fossils continues to inspire fresh theories, debates, and discussions about the evolution of

bipedalism (Senut et al., 2017:286). This dynamic nature of scientific inquiry means that our

understanding of bipedalism's evolutionary path is continually evolving, and discoveries may

challenge and existing theories. Nonetheless, the development of bipedalism remains a crucial

factor in shaping human history and distinguishing us from our ape ancestors. It remains a

symbol of the dynamic forces that have shaped the incredible history of Homo sapiens. (Senut et

al., 2018:286).
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