Human Evolution 2009

Bipedalism
Topics:
• Major changes in the skeleton related to
bipedalism (modern humans contrasted with
modern apes)
• Head - position of foramen magnum
• Hips - bowl shaped - ilium is short and broad;
ischium is short
• Knee - valgus angle; adduction
• Feet - great toe (hallux) adducted.

• Explanations for bipedalism

• Kingdon’s (2004) hypothesis of the “groundape”
• The switch from facultative to obligate
bipedalism.
Single footprint from Laetoli:
http://anthropology.si.edu/
humanorigins/ha/laetoli.htm
 On being a biped

Clockwise from top:

1. Meerkat, 2.American
Crow, 3. Kangaroo
Images from:
1-http://www.fellow
earthlings.org/
2,3 Animal Diversity
Web -
http://animaldiversity.
ummz.umich.edu/site/
index.html Dinosaur reconstructions © OUMNH Gareth Monger.
http://www.dinosaursociety.com
 Features of the modern human skeleton which are
related to bipedalism
Ilium

Ischium

In apes the pelvis is long and narrow, and

has a box-like shape. In humans it is short
and broad and is bowl shaped.
The ilium is shorter and broader in humans -
this is the “hip-bone”.
The ischium is also shorter in humans – this
is the part of the hip surrounding the
acetabulum where the femur articulates

Lewin & Foley (2003) Figure 9.13

Lewin & Foley (2003) Figure 9.11

 Valgus Angle

Valgus angle

In humans and other bipedal hominins the knee is angled in In apes it is not.
towards the midline of the body (adducted), bringing it
under the centre of gravity.
Lewin & Foley (2003) Fig 9.12
 The position of the foramen magnum

Because the skull is perched on top of a vertical spine in a biped, the foramen magnum
- the “large hole” through which the spinal cord enters the cranium – is located towards
the centre of the skull. In apes it is further back. However, you also need to remember
that apes are characterised by an orthograde posture, and in a typical quadruped you
would not see the foramen magnum as a hole on the underside of the skull at all.

Lewin & Foley (2003) Fig 9.14

 The unusual ape

Modern African
ape Modern
human
Lewin & Foley (2003) Figure 8.13
 Biomechanics of walking

By being able to balance on a fully extended, straight

leg during the stance phase, humans reduce the
energetic cost of walking.

The figures above and on the left

both illustrate the same point – the
abductors linking pelvis and femur
contract to counteract the
tendency of the body to collapse
towards the unsupported side
during walking. However, as the
figure above shows the pelvis still
tilts during walking, but this is
minimised in humans compared to
Top two figures from Lewin & Foley (2003); bottom from Boyd & Silk (2003).
apes.
 Chimpanzees & bipedalism

Chimpanzees use a
variety of postures.
Their main mode of
slow locomotion on
the ground is
knuckle-walking, as
shown by the pair
in the distance in
the illustration on
the left.
Bipedal postures
and, occasionally,
brief periods of
locomotion are
seen most
frequently during
threat displays (left)
or when feeding
(right).
Figure from Boyd & Silk (2003)

Figure from Fleagle (1999)

 Ape, Australopithecine and Human compared

We can see that fossil evidence shows the Australopithecines to be more like humans
than chimps in features of their lower limb morphology that relate to bipedalism.
However, it is also clear that Australopithecines are not identical to modern humans in
these features.

Figure from Campbell, B. Ed (1979) Humankind

Emerging. Little.
 Laetoli footprints
From 1977 to 1979 a team led by
Mary Leakey (below) uncovered
the trails of bipedal individuals
which had been made in volcanic
ash falls about 3.6Mya. The
individual footprints (left) are
remarkably similar to modern
humans.
From the stride length (right) it is
possible to estimate the height of
the hominins. One was between
3’8” and 4’4” and the other
between 4’4” and 4’11”.

© John Reader

Single footprint:
www.mnh.si.edu/anthro/
humanorigins/ha/laetoli.htm

Mary Leakey:
www.leakeyfoundation.org Trail: http://www.liv.ac.uk/premog/premog-
research-current-laetoli.html
 Bipedalism

• Major changes in the skeleton related

to bipedalism (modern humans
contrasted with modern apes)

• Explanations for bipedalism

• Kingdon’s hypothesis of the “ground

ape”

•The switch from facultative to obligate Reconstruction drawing of

Australopithecus afarensis from Fleagle
bipedalism. (1999)
 Explanations for bipedalism

Some of the explanations put forward for the evolution of bipedalism.

From Fleagle (1998)
 Thermoregulatory model of bipedalism
The figure below shows that an
upright stance means that a
bipedal hominoid would absorb
60% less heat at midday than a
quadrupedal hominoid.
Lewin & Foley (2003) Figure 9.17

The diagram above illustrates how bipedal

locomotion helps an animal living in warm
climates to keep cool –
* by reducing the amount of sunlight that falls
on the body,
* by increasing the animal’s exposure to air
movements
* and by immersing it in lower temperature air.

Boyd & Silk (2003)

 Bipedalism

• Major changes in the skeleton related

to bipedalism (modern humans
contrasted with modern apes)

• Explanations for bipedalism

• Kingdon’s hypothesis of the “ground

ape”

•The switch from facultative to obligate Reconstruction drawing of

Australopithecus afarensis from Fleagle
bipedalism. (1999)
 Kingdon’s
hypothesis of the
“ground ape” – 1

Sketches of imaginary squatting ground ape, illustrating

altered balance between upper and lower trunk with an
enlarged figure of the supposed proportions when
standing

Ape proportions when squatting

(top) compared with those of a
“ground ape” (right) and a human
(bottom)
Why apes and their
descendants would benefit
Figures from Kingdon (2004) from long arms whilst
foraging on the ground
Kingdon’s hypothesis of the “ground ape” - 2

A ground ape would

have benefited from
being able to twist its
upper torso and
increase the foraging
reach before moving on

Apes have relatively

large, long pelves,
which restrict movement
at the “waist” – unlike
humans where the
reduction in the length
of the pelvis gives more
freedom for the upper
torso to twist at the
waist.
 Biogeographical considerations for evolving
bipedalism

What would tempt an ape onto the ground?

Where might such conditions exist?

Figures from Kingdon (2004)

 Bipedalism

• Major changes in the skeleton related

to bipedalism (modern humans
contrasted with modern apes)

• Explanations for bipedalism

• Kingdon’s hypothesis of the “ground

ape”

•The switch from facultative to obligate Reconstruction drawing of

Australopithecus afarensis from Fleagle
bipedalism. (1999)
 Chimpanzees & bipedalism

Chimpanzees use a
variety of postures.
Their main mode of
slow locomotion on
the ground is
knuckle-walking, as
shown by the pair
in the distance in
the illustration on
the left.
Bipedal postures
and, occasionally,
brief periods of
locomotion are
seen most
frequently during
threat displays (left)
or when feeding
(right).
Figure from Boyd & Silk (2003)

Figure from Fleagle (1999)

 Energetics of locomotion

Compared to
quadrupedal
mammals, chimps
are less efficient at
both walking and
running – they lie
above the
regression line.
Humans are less
efficient at running
but more efficient at
walking.

Lewin & Foley (2003) Figure 9.15

 Costs and benefits of
climbing and walking
The energetic costs of locomotion
are an important consideration
when trying to work out why
bipedalism may have evolved.
Whilst bipedalism is energetically
efficient on the ground, it is more
costly in the trees – thus, as
shown on the top graph - more
time spent on the ground will lead
to selection for bipedalism. The
bottom graph shows how
modelling the energetics in terms
of daily time budgets suggests
that the critical zone for transition
to bipedalism is where about 60-
70% of the time is sent on the
ground. This implies that it would
pay a hominin to be bipedal even
while it is still spending a lot of
time feeding in the trees.
Lewin & Foley (2003) Figure 9.18
 Habitat change & facultative
bipedalism

In the latter part of the Miocene the forests start

to break up because of climatic drying. As shown
above, this means that the daily range of an
ancestral hominin would no longer be
encompassed by continuous forest. The two
pictures on the right give impressions of the Afar
region of Ethiopia at the beginning of the
Pliocene, showing the clumps of woodland
interspersed in more open habitat.
Figures (clockwise from top) from Lewin & Foley (2003); Turner &
Anton (2004); Fleagle (1999).
How did the early hominins walk?
Sellars et al (2005) Have used some
clever robotics programs to simulate the
two main ways in which Lucy might have
walked.
The top sequence shows her moving with
a chimpanzee-like bent-knee, bent-hip
gait. This is energetically more costly, but
would have been stable.
The bottom sequence shows her moving
with a human-like straight leg, extended
hip gait. This is an energetically more
efficient gait, and it is also stable.
Both of these simulations could have
produced the footprint trail found at
Laetoli.
Sellars et al (2005) Journal of the Royal Society
Interface 2 431-441
Image from:
http://www.liv.ac.uk/premog/premog-about.html
 Measuring locomotor costs

A recent study by Sockol et al

(2007) compared the energetics
of locomotion for chimps moving
either quadrupedally or bipedally
with humans. By looking in detail
(below, right) at individual
performance, it is possible to see
that some individual chimps can
move more efficiently when
bipedal than when quadrupedal.
Close study of the biomechanics
of movement in these individuals
shows that a critical variable is
the extent to which they extend
the hindlimb. This should focus
attention on what can be inferred
about hindlimb extension from
fossil hominins.

Sockol et al (2007) Chimpanzee

locomotor energetics and the origin of Net cost of transport for chimpanzee quadrupedal walking
human bipedalism. PNAS 104 12265- (blue), chimpanzee bipedal walking (red), and human
12269 walking (yellow). Dashed lines indicate trendlines for
running and walking in birds and mammals.
 Variations on bipedalism
Lucy compared to a modern
human
From Boyd & Silk (2003)
Left: There are clear
contrasts between the
skeletons of an
Australopithecine – in
this case “Lucy” (A.
afarensis) and a
modern human in terms
of limb proportions and
also the shape of the
rib-cage and pelvis.
Right: It is not until we
get to Homo
erectus/ergaster –
illustrated here by the
nearly complete
Nariokotome boy – that
we see a skeleton
which is “modern” in
terms of the proportions
of its postcranium.
Nariokotome boy
Lewin & Foley (2003) Figure 13.8
Facultative & obligate bipedalism
The skeletons of Homo ergaster
(African H. erectus) and Au.
afarensis compared.
Enlarging Lucy’s skeleton
emphasises the greater breadth
relative to height of the
australopithecine build, as well
as showing the difference in
proportion of the arms, torso
and legs compared to the more
linear build of later Homo.
This build is better adapted for
long distance walking and
running.
This change in proportions is
also consistent with
thermoregulatory constraints on
body shape (shown by the
figures for body surface: body
mass given in brackets).

Figure from Conroy (1997)

 How important was endurance running?

Bramble and Lieberman (2004), argue that selection for endurance running has been very
important in the evolution of the “modern” human build. They focus on some of the
differences between chimps and humans (left) as indicating this and point out that
endurance running in modern humans (right) is unusual in having a flat rather than U shaped
curve for the “cost of transport” (COT) at speeds used for endurance running (this includes
the average jogging speed of approx 3.5 m/sec).
Bramble & Lieberman (2004) Nature 432, 345-352 – information & figure on right. Figure on left from
http://www.naturalhistorymag.com/0405/0405_biomechanics.html
 Bipedalism

• Major changes in the skeleton related

to bipedalism (modern humans
contrasted with modern apes)

• Explanations for bipedalism

• Kingdon’s hypothesis of the “ground

ape”

•The switch from facultative to obligate

bipedalism. Single footprint from Laetoli:
http://anthropology.si.edu/
humanorigins/ha/laetoli.htm