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two legs (Figure 17.8). Many skeletal changes
accompanied the switch to walking upright,
including the loss of opposable toes, as you will
notice if you try touching your little toe with the
big toe on the same foot.
The loss of opposable toes that accompanied
walking upright would have been a handicap in
trees, since opposable toes help grasp branches
during climbing. It is therefore likely that biped-
alism was an adaptation for living on the ground.
Walking on two feet freed the hands to carry
food, tools, and weapons, and it also elevated the
head, enabling the walker to see farther and over
more things.
The shift to life on the ground was probably
not sudden or complete. The skeletal structure of
some of the oldest fossil hominins (3–3.5 million
years old) indicates that they walked upright.
However, foot bones and fossilized footprints
show that the hominins living at that time still
had partially opposable big toes (Figure 17.9).
Perhaps they still occasionally climbed trees.
The earliest known hominin is Sahelanthro-
pus tchadensis, identified from a 6- to 7-million-
year-old skull discovered in 2002. Other early
hominins include Ardipithecus ramidus, who
lived 4.4 million years ago, and several Australo-
pithecus species that are 3–4.2 million years old,
including the first full-time walker with the first
modern foot: Australopithecus afarensis. All
of these hominins are thought to have walked
upright. Their brains were still relatively small
(less than 400 cubic centimeters in volume),
and their skulls and teeth were more similar
to those of other apes than to those of humans
(Figure 17.10). A typical modern human has a
brain volume of about 1,400 cubic centimeters,
about the same volume as a 1.5-liter soda bottle.
Within the hominin branch is the Homo genus.
The fossils identified at Riparo Mezzena were
believed to be Homo neanderthalensis bones, but
they had never been closely studied. So, Longo,
the curator from Verona, assembled a team of
Partially opposable
big toe
These bones
(shown in tan)
are based on
fossils of a
similar age.
These hominin
foot bones,
3–3.5 million
years old, were
discovered
in 1995.
These footprints of two early
hominins walking upright, side
by side, were found in Africa.
Figure 17.9
Early hominins had an upright stance and partially opposable
big toes
Fossilized foot bones show that some hominins living between 3.5 and
3 million years ago walked upright but had partially opposable big toes.
Q1: What other reason besides continuing to use trees might explain
why early hominins had partially opposable big toes?
Q2: In what way does the pattern of footprints in this figure suggest
that the print makers were walking upright?
Q3: Why do you think we no longer have partially opposable big toes?
Q1: Through natural selection, deleterious traits will tend to disappear from a population over time. Which traits might
have been deleterious for ground-dwelling early hominins?
Q2: Through natural selection, advantageous traits will tend to persist in a population over time. Which traits might have
been advantageous for ground-dwelling early hominins?
Q3: Adaptations to upright walking also mean that human females have more difficulty giving birth than do females of
other species. What adaptation would you predict has had the greatest impact on this result?
researchers to analyze them, including Condemi,
the anthropologist. Condemi had long been inter-
ested in the movement of Neanderthals across
Europe and how their populations overlapped
with modern Homo sapiens populations, and she
wanted to compare the Riparo Mezzena fossils to
fossils from Neanderthal groups that had been
dug up elsewhere around Europe, hoping to see
what the fossils might say about how Neander-
thals and modern humans interacted.