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Scientists are accumulating ever more data to unravel the tale of human evolution, but as
they are doing so, the story is becoming ever more complex. At every step, it has
challenged and enlarged our idea of who we are as a species. This lecture offers an
overview of the current state of our knowledge, resultant insights, and perplexing
mysteries, both old and new.
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Find at least three insights about primates (evolution, history, ecology, life
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history, behavior) from reading this map.
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Who is missing from the map?
Recall from the previous lecture that primates are thought to evolved from
small mammals akin to tree shrews.
Before you look at the next slide, try to recall what the key primate
characteristics are.
Primates evolved to be arboreal, that is, to living in trees. The earliest lineages
with distinct primate features appeared well after the Cretaceous Extinction.
Prosimians (tarsiers and lemurs) are almost entirely arboreal and share
features characteristic of early primates more than do the anthropoids.
Anthropoids include monkeys, apes, and human, and have more derived
characteristics (that is, developed from ancestral forms) of eyes, snout, and
brain. These shared features indicate that Old and New World primates likely
shared a common ancestor before the continent of South America split off from
Africa. The rift between Africa and the westward-drifting South America
opened to full separation by the widening Atlantic Ocean by around 110 mya.
Thereafter, evolution of the Old and New World primates proceeded
independently.
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This chart displays the allometric relationship between brain and body size
among primates. Notice the units on the X and Y axis! Based on this chart,
what can you say about the following:
Differences in body size between prosimians and cercopithecoids?
Cercopithecoids and hominoids?
Differences in brain mass between prosimians and hominoids?
Why is it useful to evaluate brain mass in relation to body mass?
Why is Homo sapiens considered the odd one out?
Overall, is the relationship of brain size to body size linear (one to one)? What
does this mean?
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An important fossil that shows how the lineage looked before the Old World
monkeys and apes split (red asterisk) is Aegyptopithecus.
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As you can see from the characteristics listed here, hominoids did not evolve
startlingly new features; rather, they extended evolutionary lines already
established among primates.
What does the lack of a tail suggest?
Any thoughts about why they have 5-pointed molars?
If a species exhibits pronounced sexual dimorphism in a species, what does
that suggest about its social relations and reproductive patterns?
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The divergence of the Old World monkeys and hominoid lineages occurred at
around 20 mya, as documented by the fossil Proconsul.
From this chart, date the approximate time of the last common ancestor between
cercopithecoids like the macaque, and the great apes like the gorilla.
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During the Miocene (23 5.3 mya), the apes ruled the primate world. Its been
estimated that as many as 100 species ranged across Europe,Asia and Africa. But
ultimately, as far as we know, only one of these species evolved into a creature
with unusual skills in language, stone tool making, theory of mind and cultural
learning.
How was the brain of these ancestral species modified during evolution so
that humans alone manifested these particular, not to say peculiar, mental
abilities?
Will focusing on the brain alone answer this question?
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This chart shows the approximate date of divergence (and thus of last
common ancestor) for the taxa among catarrhines. Hylobatids represent one
family, and hominids are another family. Does this answer a question posed
on slide 6:
Who is more closely related, gibbons and orangutans, or orangutans and
chimpanzees?
It may help to compare how long ago the lineages of gibbons and orangutans
diverged, with the time of divergence between orangutans and chimpanzees?
Why?
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bipedal
highly encephalized
complex technology
highly innovative
highly social
elaborate system of communication
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Summary I
Anthropoids and prosimians are the two suborders of the
Primate order. Monkeys, apes and humans are anthropoids.
Prosimians have more primitive characteristics and anthropoids have
more derived characteristics. Each suborder has several families
within it.
2.
The earliest true primates, the adapids and omomyids, appear in the
fossil record 55 mya
3.
4.
5.
6.
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The smudged red disk below the hand stencils is the oldest cave art yet dated, at
40,800 years old. Located in El Castillo cave in the Cantabria region of northern
Spain, this image might have been created by Neanderthals.
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As this chart plots, accumulating evidence makes clear that humans and apes
diverged long ago, at least 6 mya in the case of chimps and humans. All of the
creatures on the lower track indicated by the blue box are called hominins.
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What is a hominin?
A species that is more closely related to
modern humans than to modern
chimpanzees
Inferred primarily from evidence for
bipedalisnn
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The two skulls on the lower right show the view from below. Take a
closer look at insertion of the spine into the skull and the shape of the
pelvis. Again, why does bipedalism require these changes?
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Hominin Phylogeny
Orrorin
Ardipithecus
ramidus
HRH CHIMPANZEE
Au. afaren i
HOMO
Au. africanus
i
Sahelan thropus
Au. anamensis
Au. robustus
Au. aethiopicus
Au. boisei
The fossil record on early hominins is thin for the first
2 million
years .
6 MYR S M
AGO
4 MYR
AGO
3 MYR
AGO
2 MYR
AGO
1 MYR
AGO,
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Two different genera have been found that exhibit cranial features compatible
with more bipedal habits.
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The introductory lecture touched on the limitations of fossil evidence and the
rarity of finding complete specimens, so paleontologists usually are working with
bones and bone fragments. Therefore, there was tremendous excitement over the
discovery of a partial skeleton from about 4.4 mya. You can see that the toe
diverges which indicates climbing but the rigidity of the foot seems to suggest
bipedal capacity. Evidence of the ecology from the location of the remains
indicates it occupied a wooded rather than grassland habitat.
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Ardipithecus ramidus
I.
4.4 nnya
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The great majority of the early hominin fossil record discovered so far was
located in East and Southern Africa. There is a big gap between these two
areas, as you can see on the map, so there must have been hominins living in
between. Again, we need to keep in mind how partial our record of our
evolutionary past still is, although more is being discovered every year.
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Notice the presence of forms with a light build and forms with a much more
heavy build, which is thought to relate to different feeding strategies.
Australopithecines were definitely bipedal, but looking at the chart, what
would you conclude about the evolution of cranial capacity over this long time
period?
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One of the most famous fossils that shows us what early bipeds were like is
another partially complete skeleton, Lucy, who was a member of the earliest
known Australopithecine species, afarensis, that persisted for nearly a million
years. Again, the relative completeness of the skeleton allowed
paleoanthropologists to extract a huge amount of information about her.
Notice the shape of the pelvis: why is that a very telling indicator of bipedal
gait?
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Recall we said during the first lecture that behavior does not fossilize. The
Laetoli footprints were a spectacular exception to this rule: three individuals,
two adults and a child, walked together through a fresh deposit of volcanic ash.
Subsequent damp turned the ash to a solid Look at the footprint. Watch the clip
of the film linked here and listen to what Attenborough tells us about the many
lessons from these footprints. Notice how long ago this was!
Show Laetoli min 23:40-26:00
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So the great majority of early evolution in the hominin lineage relates to the
emergence and then long persistence of bipedal forms in the absence of cranial
expansion. Lets consider again the potential benefits of bipedalism that
established and maintained this mode of locomotion.
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Is bipedalism an adaptation?
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We know that brain expansion occurred along the hominin lineage, such that
brains are three to four times larger than chimpanzee brains and nearly three
times that much large than Australopithecines. A major question about human
evolution is how and why this occurred. At the outset, paleoanthropologists
expected that cranial expansion would co-evolve with bipedalism, freeing of
hands, and so forth.
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Direct evidence of human brain evolution comes from the very rare fossilized
skulls of our ancestors. These skulls can be used to make endocasts, as
illustrated here. Endocast volume can then be measured as an approximation of
brain volume.
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Over the course of human evolution, brain size tripled so that the modern
human brain is the largest and most complex of any living primate. The
emerging answers to when (and why) this happened are surprising.
Endocasts of H. erectus, left, and H. sapiens, right.
From: http://humanorigins.si.edu/human-characteristics/brains
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This graph plots braincase volume (thus, presumably, brain size) over time,
making it clear that notable cranial expansion did not begin until about 2 million
years ago. Once it began, however, brain size increased consistently over time to
achieve current cranial capacity relatively late in human evolution.
Can you think of selection pressures that might have favored bigger brains and
thus could account for this pattern?
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Hominid Phylogeny
HONCH
O
CHIMPANZE
E
HOMO
Au. anamensis
Au. aethiopicus
Au. boisei
6 MYR
AGO
5 MYR
AGO
4 MYR
AGO
3 MYR
AGO
1
MYR
AG O ,
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e
n
Enlarged brain
Australopithecus africanus
4413cc
Homo erectus
101110cc
Homo sapiens
1300 + cc
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This chart shows you brain size by species over time. Cranial expansion began
with Homo habilis and continued through the history of the genus Homo.
Note when body size increased.
What does that suggest?
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Note the behavioral, physical, and ecological correlates of brain size over
time. These begin to tell us more about selective pressures.
Brain size increases slowly from 62 million years ago
During this time period, early humans began to walk upright and make simple
tools. Brain size increased, but only slightly.
Brain and body size increase from 2 million800,000 years ago
During this time period early humans spread around the globe, encountering
many new environments on different continents. These challenges, along with an
increase in body size, led to an increase in brain size.
Brain size increases rapidly from 800,000200,000 years ago
Endocasts of H. erectus, left, and H. sapiens, right.
From: http://humanorigins.si.edu/human-characteristics/brains
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We plot changes in behavior and ecology (blue boxes) against the pattern of
changing brain size.
For a long time, tool use was considered a mark of intelligence that would open
new dietary niches and life history strategies. Thence, the expectation was tool
production would accompany increased brain size because tool use.
From this chart, would you say that the evidence supports that expectation?
What does this suggest about hominin evolution?
Notice that geographic expansion of the species outside Africa apparently
occurred only with the start of cranial expansion, among the genus Homo.
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Do you remember Rick Potts idea about the ecological stimulus for
brain expansion? Review this segment in the PBS program that you
watched.
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Human brain size evolved most rapidly during a time of dramatic climate
instability and change.
Why might the onset of climate instability and opening of novel environments
have created selective pressures favoring expanding brain size?
From: http://humanorigins.si.edu/human-characteristics/brains
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Current thinking suggests that larger, more complex brains enabled early
humans of this time period to interact with each other and with their
surroundings in new and different ways. As the environment became more
unpredictable, bigger brains helped our ancestors survive.
This skull is a representative of the first species in our genus Homo.
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Production and use of complex tools also is considered a key feature of the
human lineage. The first known tools, known as Oldowan, are dated at about
2.6 mya, and were associated with late australopithecines. The photos show
examples. Upper right is a chopper, lower right are flakes and the core they
were struck from. And these are the tools associated with Homo habilis.
Think about what you can do with tools like these.
If you are interested in early tool use and its relationship to human evolution,
especially brain evolution, we have a world expert on the Anthropology faculty
here, Prof. Dietrich Stout.
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The maps shows the distribution of bones at an ancient butchery site.The bone
fragment comes from that butchery side. Look at the places on the bone indicated
by the arrows.
Here is exciting evidence that the tools were used to butcher carcasses. The
arrows point to cut marks on the bone clearly made by a sharp edge rather than a
tooth.
If you are interested in early human behavior, tool use, and ecology, we also
have a world expert on the Anthropology faculty, Prof. Jessica Thompson.
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Watch carefully the segment of this program showing a kudu hunt. What skills
are required to perform this? We will discuss this further in future lectures, but
this will get you started thinking about it. From what you have seen about the
record of human evolution in this lecture, you will know that the vast majority
of human evolution (for at least 2 million years since tool use became quite
common) was spent in the forager, hunting-gathering niche.
Hunting: 31:29 35:44
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Changes that distinguished the next member of the lineage, Homo erectus, showed
some increase in brain volume but a substantial increase in height. This species
was extant for over a million years, and was the first member of our genus to
migrate outside Africa.
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Here you can compare two famous, relatively complete skeleton finds,
Australopithecus africanus Lucy and Homo erectus Nariokotome Boy.
See how many differences you can find between these two.
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This is a representative Homo erectus skull. Compared to those you have seen
previously, what are distinctive features here?
Note height and angle of forehead, brows, cheekbones, plane of lower face and
angle of teeth. Use of tools and fire for processing foods reduced the need for
powerful jaws and heavy teeth, permitting a reduced muscles around face and jaw,
lighter cheekbones and more acute angle of jaw for chewing rather than pulling
and tearing.
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Acheulean tools are worked symmetrically on both sides. Such tools are
characteristic of Homo erectus and first appeared when the species appeared.
This tradition persisted for over 1.5 million years, from ~1.8 to ~0.1 mya.
It may seem rather dull to maintain a single tradition for such an immense
length of time (think of how rapidly our technology changes), but consider
what is required to maintain such fidelity over that long period. Moreover, the
method of preparation was applied to produce an increasingly wide array of
tool types.
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The oldest known evidence of hominins outside of Africa are Homo erectus
fossils that come from the site of Dmanisi in the Republic of Georgia, one of the
most prolific fossil human sites in recent years. The age of the
Dmanisi fossils is about 1.85 to 1.75 million years old. Comparing the
Georgian and Chinese dates, the evidence from the Nihewan and Yuanmou
regions is consistent with the spread of early hominin populations beyond
Africa into the Caucasus region and into East Asia between roughly 2 million
and 1.7 million years ago.
By 1.66 million years ago, early humans of the genus Homo who reached
eastern Asia were able to disperse over a wide area that extended from at least
40N (Nihewan basin) to 7S (Java, Indonesia), across a habitat range from
temperate grassland to tropical woodland and possibly forest.
From: http://humanorigins.si.edu/research/asian-research/earliest-humanschina
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As more finds are made, the hominin record is getting more and
more complicated. This chart on the upper right shows that:
The map on the left shows hominin sites, documenting the wide
distribution of Homo erectus and later forms.
Not included on either graph is the diminutive Homo floresiensis,
found in Indonesia in 2003, living ~93000 13000 ya.
Who knows, there may be other species of early Homo that have not
been discovered.
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More fossil evidence from diverse sites across Africa, Europe and Asia plus the
ability to analyze ancient DNA has greatly increased both the complexity of the
record and our ability to discern the tangled, evolving web of Homo species
through time. We now clearly know that human evolution emphatically was not
a neat, linear story that lead directly to our species. Rather, many species have
been woven into the history of an evolving lineage that both evolved diverse
forms yet also maintained linkages through interbreeding.
H. sapiens skull is Cro Magnon, early modern humans in Europe earliest dated at
43000 ya
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Neanderthal Range
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The map above shows the likely paths of human migration out from the
location in Africa where modern humans probably first appeared. The alreadyoccupied range of Homo erectus is shown for comparison. (Not shown is the
radiation of Homo heidelbergensis into Europe.) Also, that of Homo
neanderthalensis.
https://en.wikipedia.org/wiki/Acheulean#/media/
File:Spreading_homo_sapiens_la.svg
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Hominidae: living humans, all human ancestors, the many extinct members of
Australopithecus, and our closest primate relatives, the chimpanzee and gorilla.
According to The Tree of Life by Guillaume Lecointre and Herv Le Guyader
(Harvard University Press: 2006), the similarly named and easily confused
categories of humans and near human apes, in order of increasing inclusiveness,
are:
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By now you probably have a good sense of the answer to the question of why
our knowledge of human evolution remains incomplete. Indeed the full story
likely will never be known, but we are learning a great deal not only about
ourselves and our assumptions, but also about evolution and ecology in
general.
Homoplasy is the sharing of a common trait among related species whose
common ancestral species did not have that trait. Homoplasy reflects
convergent evolution as species adapting to the same niche evolve
characteristics to meet the selective pressures of that niche.
Why might the search for the Last Common Ancestor with our ape relatives (or
any other species) be a fruitless quest? For one, the LCA surely was a
population. For another, that population also would have been comprised of
many traits neither chimp nor human-like. Both species evolved tremendously.
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Summary II
Species that are more closely related to modern humans than
modern chimpanzees are known as hominins
Hominins are identified by skeletal specializations for bipedalism and by reduced canine
teeth
2.
3.
4.
The evolution of the genus Homo involved a dramatic increase in
brain size and a reduction in the face, jaws and teeth
Neanderthals had large brains but were outcompeted by Cro-Magnons (anatomically modern
humans in Europe)
All living humans are descended from a small population that lived in
Africa approximately 100-200,000 years ago. Modern and archaic humans
interbred.
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