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feet. These anatomical adaptations evolved over millions of years and differences exist between
earlier and later hominin species (Jurmain, Kilgore and Trevathan). Australopith and
paranthropine evolution show a notable step in the evolution of humans because these species are
among the earliest hominins and are known to have evolved the adaptation of bipedalism. The
most famous fossil of Australopithecus afarenis is the skeleton known as Lucy. From the fossil
record we can determine approximately when these traits evolved so that humans could become
bipedal (Schwartz and I.).
Hominin
Date Range
Sahelanthropus
6 - 7 Ma
A. anamensis
4.2 - 3.9 ma
A. afarensis
3.9 - 2.8 Ma
A. africanus
2.8 - 2.2 Ma
P. aethiopicus
2.5 - 2.3 Ma
P. robustus
1.8 - 1.0 Ma
P. boisei
(Hominid Species)
Lets take a closer look at the characteristics of bipedal hominids. The placement of the
foramen magnum (the large hole in the cranium where the spinal cord attaches) is different from
our quadrapedal ancestor. In a quadrapedal ancestor, the spinal column also runs parallel to the
ground so the foramen magnum is located toward the back of the cranium. In a bipedal organism,
the spinal column runs perpendicular to the mandible and the ground.
Maintaining balance is crucial to walking on two legs. When humans walk they must
balance on one leg while lifting the other off the ground and swinging it forward. The center of
gravity is particularly important for this to happen. The center of gravity for quadrapedal
hominins is located near the center torso. In modern Humans who are bipedal, it is located
closer to the center of the pelvis. As the legs alternate the center of gravity shifts from one side to
the other. The lumbar curvature of the spine helps to bring the center of gravity closer to the
bodys midline and above the feet. The size and number of the lumbar vertebrate are different
than in apes. Humans have 5 larger lumbar vertebrae. Most large apes have 4 that are relatively
smaller than those of humans. The bigger the number and size of the vertebrae forms a back that
is more flexible and permits the hips and trunk to swivel forward when walking.
Australopithecus lumbar vertebral bodies were broad for effective weight transmission from the
upper body to the pelvis. Australopiths had 5 or 6 lumbar vertebrae that articulated to form a
distinctive lumbar curvature, similar to the morphology of modern humans. Another difference
with the spine is the Sacrum. The shape of the sacroiliac joint is a reflection of the lumbar curve.
The sacrum is relatively broad in modern humans with large sacroiliac joint surfaces. Modern
chimpanzees have a relatively smaller sacroiliac joint surface. These size differences are related
to the different patterns of weight distribution through the pelvis during quadrapedal and bipedal
locomotion. The australopith sacrum has a relatively large, but less curved sacroiliac joint than
that seen in modern humans (Stern).
The modern pelvis has relatively larger hip joints and larger pelvic outlet relative to
australopiths. In bipedals, the hips support and balance the weight of the torso during
locomotion. However, as the size of the pelvic outlet increased, the hip joints were repositioned
relatively further from the center line of the body. As a result, more force is exerted on the hip
joint as the joint (acetabulum and femoral head) moves further away from the bodys center of
gravity, and thus affects stability as the weight of the torso presses downward toward the middle
of the body. This issue is resolved through several adaptations in the pelvis and femur. In the
pelvis, an enlarged hip joint allows more stress to be absorbed and accommodates a larger
femoral head.
Most quadrupedal and arboreal primates have either longer arms relative to their legs, or
arms and legs of equal length. Most humans have relatively longer legs than arms. Based on this
information, it is possible to estimate the positional behavior of a species by calculating the
humerofemoral index. This index is the length of the humerus divided by the length of the femur,
multiplied by 100: humerus length x 100/femur length.
These results calculate the overall body proportion of an organism which can then be
compared to others. The higher the index value, the longer the arms, equals the likelihood that a
primate was arboreal. Arboreal primates have ratios close to 100. For example, the mean ration
for the common chimpanzee is 97.8. Humans have a lower ratio at approximately 71.8. The ratio
of the famous A. afarensis Lucy is intermediate between modern humans and chimpanzees at
84.6.
The entire weight of the upper body is transferred through the legs and into the feet when
a bipedal is standing or walking. The femur is one of the most critical links between the pelvis,
spinal column and lower legs. The femoral shaft is generally straight, ending in two bulbous
condyles. These condyles are larger and more elliptical in bipedals when compared to the
relatively smaller and rounder condyles of a quadrapedal animal. The distal end of the fermur
forms a joint with the lower leg and the knee cap at the knee joint. Another adaptation is the long
femoral neck. In humans, hips are wide apart, the shaft of the femur is angled so that the knee is
closer to the bodys midline than the hips. This angle is called the bicondylar angle, and the
resulting knee joint is referred to as a valgus knee. The object of this trait is to bring the knees
closer together, placing the feet directly below the center of gravity. One of the results of the
bicondylar angle pulling the knees together is that the tibia stands almost parallel with the bodys
center of gravity. The tibia is also relatively larger and elongated compared to quadrapedal
animals.
The distal end of the tibia forms a joint with the talus at the ankle. Human feet have also
evolved. The foot is the platform that supports the entire weight of the body including all the
forces generated by walking, running, and jumping. All of this weight passes through only one
foot at a time. To accommodate this, the foot evolved. The Humans have the most distinctive feet
of all the apes. The big toe in humans is much larger and sturdy in construction than the other
four toes. It is often referred to as a non-opposable big toe because the hallux is restricted to an
adducted position which means that the big toe runs parallel to the other toes and lateral
movement is limited (Harcourt-Smith and Aiello). In general humans toes are shorter in length
than other primates. Humans also have almost no grasping ability in their toes and feet like other
primates do. The heal bone is also relatively large in humans compared to chimpanzees
especially the back portion known as the calcaneus tuberosity. The calaneus tuberosity provides
stability and helps to absorb high forces encountered during heel strike. The shape provides
attachment points for strong ligaments that run from the arch of the foot to the tibia. The
ligaments add support and create a double arch system that helps to absorb the stress of the foot
hitting the ground. Evidence from the Laetoli Tracks in Tanzania, where footprints from several
australopiths were preserved in volcanic ash, indicates that Australopithecus had relatively short
toes, and an intermediately adducted hallux. Fingers have also changed. The degree of curvature
in the phalangal shaft coincides with the frequency of arboreal behavior (Alexander). Primate
who spend a lot of time swinging in the trees, grasping, or suspending from curved branches
have dramatically curved fingers and toes which allows for a better grip. Non-arboreal primates
like humans have relatively flat manual and pedal phalanges because they did not spend as much
time in the trees which has facilitated the evolution of precise hand movements necessary for
making tools. Australopith phalanges are intermediately curved between those of modern
humans and great apes, suggesting that climbing and arboreal behavior continued to play some
role in the lifestyle of these early hominins.
Early researchers hypothesized that large brain size was the first hallmark of the hominin
lineage. Beginning in the mid 1800's until the early 1900's, almost all known fossil hominins
had relatively large brains. The large brain bypothesis was falsified after the discovery of early
hominin fossils exhibiting ape-sized brains and bipedally-adapted morphology.
In 1924, Raymond Dart identified the fist australopith fossil, known as the Taung Child,
from South Africa. This fossil belonged to the species Au. africanus and had a relatively small
brain similar to the size of a modern chimpanzees. The inferior placement of the foramen
magnum, suggested that the Taung Child was bipedal. Dar's hypothesis that bipedalism evolved
before larger brains ran counter to the scientific consensus at the time. Because of his small
sample size and the fragmentary remains, debate about the timing of bipedalism and brain size
continued for the next 50 years. Until in 1974 when Donald Johanson found the nearly complete
fossilized skeleton of Lucy, a member of the species Au. afarensis dating to 3.2 Ma. Lucy was
unique at that time because she was one of the first fossils to exhibit both small relative brain
size and the highly derived features characteristic of bipedalism. As other contemporaneous and
older fossils are found, scientists continue to revise the bipedalism timeline (Hass). Today, there
is no doubt that the evidence demonstrates that bipedalism was one of the first hallmarks of the
hominin lineage, which could have led to more advances (Wood and N.). Like freeing the hands
to allow for the production of more technologically and advanced stone tools. In turn, the
production of more complex tools may have led to a higher protein diet that affected brain size.
Brewster 8
Works Cited
Alexander, R. Bipedal Animals and Their Differences From Humans. Vol. 204.5.
Journal of Anatomy, 2004. Print.
Harcourt-Smith, W.E.H. and L.C. Aiello. "Fossils Feet and The Evolution of Human
Bipedal Locomotion." Journal of Anatomy 2004: 403-416.
Hass, Randall Jr. Laetoli Footprints Preserve Earliest Direct Evidence Of Human
Human Like Bipedal Human Biomechanics. Plos ONE 5.3, 2010. print.
Hominid Species. n.d. Web. 10 4 2016.
<http://www.talkorigins.org/faqs/homs/species.html>.
Jurmain, Robert, Lynn Kilgore and Wenda Trevathan. Human Origins Evolution and
Diversity. Mason: Cengage Learning, 2013. Print.
Schwartz, JH and Tattersal I. "The Human Fossil Record: Craniodential morphology of
early Hominids." 2005. print.
Stern, J. "Climbing to the Top." Evolutionary Anthropology 1983: 113-133. Print.
Wood, B and Lonergan N. "The Hominin Fossil Record: Taxa, grades, clades." J. Ant
(2008): 354-376. Print.