Background
Information
There
is probably no other topic in evolution that is as controversial in
the public’s mind as human evolution. However, it is consistently
the topic that students find the most interesting. After all, it is
about something intrinsically interesting to us: ourselves. It is the
story of how we got here. Before we dive into what evolutionary biology
can tell us about human origins, it is important to keep one thing in
mind: understanding human evolutionary origins is just one part of understanding
what it is to be human.
Evolutionary
biology indeed has much to say about the human condition, but so do
anthropology and sociology, psychology and history, philosophy and the
arts. People are too complex to be understood from the narrow perspective
of biology or of any other single way of knowing.
—Douglas Futuyma, 1998
Overview
How
do we know what we know in paleoanthropology (the study of human
evolution)? How can we tell if a particular species walked upright or
lived in trees? Or if it lived in grassy savannas or tropical forests?
How can we know anything about the social structure of extinct populations?
As with any historical science, clues to the past are left,
and the job of the paleoanthropologist is to find these clues and to
interpret them. These clues come in many different forms; fossils of
the bones and skulls of humans are the most dramatic and well publicized.
The size of the skull can obviously tell us about the size of the brain.
In addition, the size, shape and orientation of other bones, as well
as the size of the muscle attachments on the bones tell us a lot about,
for example, how the individual moved and used its hands. Other important
clues about the past are tools and other archaeological items such as
cave paintings and jewelry; footprints preserved in the ground; animal
and plant fossils found at the same site; butchery marks found on animal
bones. In addition, genetic data from modern humans and even some fossil
specimens, can tell us about how groups are related and how long ago
they are likely to have diverged from each other.
The archaeological record of hominids1
is in many ways amazingly complete, showing many transitions between
species. Yet in other ways, it is entirely inadequate to fully answer
all of the questions we have about human origins. Some questions may
never be answered to our complete satisfaction. The archaeological record
does, however, allow us to make several statements with extremely high
confidence. First, modern humans evolved from apelike ancestors,
through a series of intermediate steps. Humans and the great apes—chimpanzees
in particular—diverged from a common ancestor as recently as about
7 – 8 million years ago (Mya); the transition from this common
ancestor to modern humans has been well documented. Second, hominid
evolution has been mosaic—important features have evolved
at different times, some sooner than others, and at different rates.
For example, large brains and bipedalism (the ability to walk upright)
are both hallmarks of human evolution; however, bipedalism evolved several
million years sooner than increased brain size.
There
have been many different transitions in hominid evolution. Here are
some of the important ones:
1)
hominids diverged from the great apes, ~7 – 8 Mya
2) bipedalism, at least 4 Mya (probably much earlier)
3) simple stone tools, ~2.5 Mya
4) more sophisticated tools (hand axes), ~1.5 Mya
5) fire mastered, spears and other tools, ~0.5 Mya
6) symbols and signs of abstract thought (jewelry, paintings, ritualized
burials), starting ~40,000 ya
7) agriculture, ~10,000 ya
Our Primate Origins
In
The Descent of Man, Charles Darwin summarized the mass of evidence
for our common ancestry with the apes. As there had been no hominid
fossils discovered at the time, his evidence came from embryological,
physiological, anatomical and behavioral comparisons of modern humans
with vertebrates (including the apes).
For
example, humans and apes have a similar skeletal structure, and they
go through the same embryonic stages (it is only late in development
that they differ, with different body parts growing at slightly different
rates). From these data, Darwin concluded that humans are vertebrates,
and more specifically primates, and that we share a common ancestor
with the apes. Since we were more similar to the African apes (gorillas
and chimps) than to the Asian apes (orangutans), he also predicted that
Africa was the place where humans originated.
Today, we have access to a tremendous breadth of accumulated evidence,
and all of it supports both of Darwin’s predictions. We now know,
for example, that modern humans share a number of characteristics with
the modern apes. Both groups have shoulders with a wide range of movement,
dexterous hands with fingers that are capable of strong grasping, nails
instead of claws, relatively large brains and greater cognitive abilities
than other animals, and large, complex social groups. These characteristics
are shared between humans and apes, and are older than the split between
these groups (shared primitive traits). Since humans and apes
diverged (about 7 – 8 Mya), both groups have evolved unique characteristics
that distinguish and that define their particular groups (derived
traits). Humans, for example, have evolved the ability to walk
upright (bipedalism), a much larger and more complex brain, a much greater
intellectual capacity, language (spoken and written), and the ability
to precisely manipulate very small objects with their hands, among others
traits. Likewise, chimps also have evolved new traits of their own during
the time from the split with hominids and the present.
Perhaps the strongest evidence that we are related to the great apes
comes from studies of our DNA. Today we know that humans and chimpanzees
are genetically extremely similar, sharing 98% of the same genes. More
specifically, the base pair sequences of our DNA are 98% identical to
the sequences of chimp DNA. Modern humans and chimps are sister species,
each being more closely related to each other than to any other extant
species (Figure 1). Chimps are actually more closely related to humans
than they are to gorillas!
Darwin
was also right about Africa being the place of origin of humans. We
now have a multitude of fossils documenting early lineages of hominids,
and all of the oldest hominid fossils have indeed been found in Africa.
Africa is the place were humanity originated.
The
First Hominids
According
to DNA comparisons, humans are thought to have diverged from the apes
~ 7 – 8 Mya. Until recently, however, the oldest hominid fossil
ever found was about 4.4 million years old (Myo): Ardipithecus ramidus.
Skeletal evidence indicates this species was bipedal, able to walk upright
on two legs. Recently (2000), a 6 Myo fossil, Orrorin tugenensis,
was found and classified as a hominid: it had small teeth, like later
hominids, with hard enamel on their cheek teeth (indicating a diet of
tough plants, rather than the soft fruit diet of chimps). There is also
some skeletal evidence (e.g. where the muscles attach to the leg bones)
that indicate this species was also bipedal. Even more recently (July
2002), an even older fossil hominid, close to 7 Myo, was found, and
is different enough from the other two ancient species to be classified
in its own genera: Sahelanthropus tchadensis, nicknamed “Toumai.”
S. tchadensis is thought to have lived closed to the time of
the divergence between chimps and humans. This fossil is further support
of the close evolutionary relationship between these humans and chimps:
it has a mix of characters that are associated with early chimps and
early hominids. There is even some evidence (the way the spine connects
to the skull) that this species may have been bipedal, although this
cannot be concluded with confidence at this time.
Bipedalism is a distinguishing characteristic of humans
that is thought to be one of the traits that evolved very early in the
human lineage. In fact, it is thought to have been a critical step in
the evolution of hominids, and it entails a suite of skeletal changes.
Compared to non-bipedal organisms, in bipeds the pelvis is more bowl-shaped
(to help support the internal organs), the upper legs are angled inward
from the hip joints (which puts the knees in a better position to support
the body), the foot is shorter with more rigid toes (to help push off),
the spine is shortened and S-shaped (for rigidity and balance), and
the opening of the bottom of the skull is more anterior (to set the
head more directly over the spine). All of these skeletal adjustments
are part of the evolutionary modifications that gradually evolved in
our ancient ancestors as they became increasingly upright in stature.
What may have led to the evolution of bipedalism? In other
words, what kind of advantage did individuals who were more bipedal
have over those who were less bipedal? Over the years, there has been
a lot of speculation about this question, and for a long time, the “savanna”
hypothesis was in favor. This hypothesis stated that the earliest hominids
evolved during a period of climate change, on the newly formed African
savannas (tall grasslands). The ability to stand upright was thought
to be advantageous because it allowed them to look over the tall grass,
presumably to detect predators. However, it appears that the earliest
hominids did not live in savannas (which are now thought to have not
been common in Africa until much more recently), but rather in forested
environments. Other possible advantages of bipedalism include: it allowed
individuals to carry food back to their family group, it enabled individuals
to walk for long distances more efficiently, it made it easier for individuals
to reach food from low branches, and standing upright decreased the
amount of skin exposed to the hot African sun. At present, the existing
data are inadequate to convincingly discriminate among these alternatives.
The
Australopiths
The first species that was undoubtedly bipedal was Australopithecus
afarensis. One of the reasons for this high level of confidence
(besides many skeletal characteristics) is the Laetoli footprints, made
by an A. afarensis individual 3.6 Myo. A. afarensis
is a well-documented species, with hundreds of fossils, including one
of the most famous hominid fossils: “Lucy,” an incredibly
complete skeleton of a young female who lived 3.2 Mya. A. afarensis
is thought to have lived from ~3.9 – 3 Mya, and is probably an
ancestor to the later australopiths and maybe even to the genus Homo.
Members of this species still had a relatively small brain (~400 cubic
centimeters, roughly the same size as modern apes), and long powerful
arms, suggesting that although they were definitely bipedal, they also
were skilled at climbing trees.
The australopiths
are commonly divided into two major groups: the early or “gracile”
australopiths, which arose more than 3 Myo, and the late or “robust”
australopiths, which arose after 3 Myo. The graciles had smaller teeth
and jaws, while the robusts had much larger faces, with large jaws and
teeth. Ardipithecus ramidus and Australopithecus afarensis
are both gracile australopiths, as are a later species: Australopithecus
africanus, which lived ~3 – 2 Mya. This species had a slightly
larger brain (~500 cc), a more rounded skull, and more modern-looking
face and teeth than A. afarensis, suggesting that it may have been the
ancestor to the genus Homo. Whether A. ararensis or
A. africanus was ancestral to Homo has yet to be determined.
The robust australopiths are a separate lineage from that which eventually
evolved into Homo. The robusts had evolved by 2.7 Mya, and
were extinct by ~1 Mya. They are referred to as “robust”
because of their large face, broad thick cheek teeth, and prominent
sagittal crest (a raised area of bone along the top of the skull, used
for jaw muscle attachment) all indicating that they were very strong
chewers, probably of tough, fibrous plants. The robusts are usually
separated into their own genus, and three species are generally recognized:
Paranthropus aethiopicus (the earliest), Paranthropus robustus,
and Paranthropus boisei.
The
Genus Homo
Members
of our own genus, Homo, made their first appearance ~2.5 Mya.
The key innovations that distinguish members of the genus Homo
are considered to be: 1) a gradual increase in brain size
(from a cranial capacity of ~400 – 500 cc in the australopith
ancestors of Homo to an average of 1350 cc in modern humans),
and 2) the advent of toolmaking (the first stone tools
appeared 2.5 Mya). Some scientists think that a change in the climate
of Africa during this time period from hot and wet to fluctuations between
dry and wet environments may have given toolmaking individuals an advantage
in obtaining alternative food (underground tubers and roots, as well
as meat obtained from scavenging and hunting), as their old food (vegetation)
became more unpredictable.
Why did brain size increase? In other words, what factor selected
for increased intelligence? It is possible that the advantage of being
able to make tools selected for increased intelligence, and therefore,
increased brain size. Or it is possible that increased sociality (larger
group size) selected for increased intelligence (the larger the group
size the more intelligence it takes to keep track of alliances, deceptions,
etc.). In this scenario, the consequent increased intelligence led to
the advent of tool-making. Whatever the reason, selection for greater
intelligence resulted in larger brain size, and a suite of other traits
as well: a larger birth canal; since brain development continues after
birth, a prolonged period of time between birth and maturity, and a
longer period of parental care of infants. These traits may have then
led to changes in social structure as well (e.g. a greater need for
more extended parental care may have led to a sharper division of labor
between males and females).
Homo habilis (“tool man”) lived between ~2.5 –
1.6 Mya, at the same time as A. africanus and the later robust
australopiths, and resembled the australopiths in their dentition. However,
the brains of individuals of this species (~650 cc) were larger than
those of the australopiths. Members of this species had hands capable
of producing tools, and some of the oldest tools have been found with
fossil H. habilis, suggesting this was the first species to
make simple stone tools. The first stone tools were made by banging
rocks together to chip off the edges, creating a simple blade, which
could chop or scrape.
H.
habilis is intermediate in many characteristics between
A. africanus and Homo
erectus2,suggesting
a transition between these species. H. erectus (1.6 –
0.3 Mya) had a
cranial capacity of ~900 cc (which increased over the lifetime of the
species), made more sophisticated stone tools (called handaxes), was
the first to master fire (by 0.5 Mya), and was the first species to
leave Africa and spread throughout Asia and Europe (between 1.7 –
1.0 Mya).
The
Origin of Modern Humans
There
are two species of late Homo: Homo neanderthalensis,
and our own species, Homo sapiens. H. neanderthalensis
lived from about 250,000 – 30,00 years
ago, and actually had a
larger brain (~1500 cc) than H. sapiens (~1350 cc), although
this may just be due to an overall larger body size. They most likely
evolved from H. erectus populations in Europe, and as they
lived in the cold northern climates of western Europe, they had many
facial and skeletal characteristics that were probably adaptations to
cold: powerful, compact bodies that helped conserve heat and withstand
cold temperatures. They also had protruding mid faces, low foreheads,
prominent brow ridges, and weak chins. Neanderthals produced relatively
sophisticated stone blades and spears.
H. sapiens, or anatomically modern humans, is a relatively
new species, with the first fossils appearing ~130,000 years ago. Modern
humans have a much smaller brow ridge, a roundedbraincase, high foreheads
(resulting from the frontal lobe of the brain positioned directly over
the face), prominent chins, and a flat face. Modern humans also have
a much lighter bone structure than Neanderthals.
H. sapiens invented tools that required a much higher degree
of sophistication and skill than any other species, including spears
that were
much more efficient to use and that put the person
using it in less danger from the animal than Neanderthal spears. They
also used new types of materials, such as bones and antlers. Starting
about 40,000 years ago, ornamentation, such as jewelry, began to appear
in the archaeological record, as did other fine artwork, such as cave
paintings, ivory carvings, clay figurines, and musical instruments.
These signaled a big change in H. sapiens society: new ways
of thinking—symbolic and abstract thought. Now, at least some
people had the time to spend on skills that had nothing to do with basic
survival.
Where and when did modern humans evolve?
There are two hypotheses that attempt to explain where modern humans
originated. The “out of Africa” hypothesis
(also called the “replacement” hypothesis) states that about
200,000 years ago, modern humans originated in a small region in Africa
from a population of H. erectus that had been isolated from
the main population. Subsequently, the new species H. sapiens
migrated out of Africa into Asia and Europe. Eventually, H. sapiens
replaced other species of hominids (H. neanderthalensis in
Europe and H. erectus in Africa and Asia).
The “multiregional” hypothesis (also called
the “continuity” hypothesis), on the other hand, states
that the evolution of modern humans began when H. erectus moved
out of Africa ~1 – 2 Mya and spread throughout Asia and Europe.
Populations of H. erectus evolved in response to the new situations
that they encountered (leading to regional differences among populations).
Gene flow between neighboring regions spread novel H. sapiens
traits throughout the entire geographic range.
Both of these hypotheses make predictions about the genetic differences
we should find among contemporary humans from different geographical
regions. The multiregional hypothesis predicts that there should currently
exist genetic differences between people from different geographic regions
that developed more than 1 Mya, while the out of Africa hypothesis predicts
that these differences should have evolved much sooner (within 200,000
years ago). Molecular studies of mitochondrial DNA (mtDNA) have yielded
an estimate of the divergence time of all human populations: ~200,000
years. This means that all contemporary human mtDNA originated approximately
200,000 years ago; this supports the out of Africa hypothesis that humans
originated in Africa. These studies have also shown that contemporary
African populations have a greater diversity in their mtDNA than do
other geographical populations. This is expected if humans originated
in Africa and then spread gradually: as new populations were founded,
they were likely subject to population bottlenecks, and during these
events they would lose some genetic variation (through the process of
genetic drift). Today, then, we would see less variation within each
of these populations than we would within the ancestors of the original
source population.
In another study, mtDNA was extracted from a Neanderthal fossil and
compared with contemporary human mtDNA. This study revealed that Neanderthals
and modern humans are not very close genetically, with the estimated
divergence time about 600,000 years ago. This also supports the out
of Africa hypothesis. Overall, the genetic
evidence3 tends to favor the out of Africa hypothesis.
Fossil evidence also tends to favor the out of Africa hypothesis. The
oldest anatomically modern human fossils have been found in Africa,
and are ~130,000 years old. The next oldest come from the Near East
(~90,000 years old). Modern humans are not seen in Europe until about
40,000 years ago (Cro-Magnon people). This chronological pattern would
be expected with a migration pattern predicted by the out of Africa
hypothesis.
On the other hand, fossil H. erectus and anatomically modern
humans in some regions (e.g. Asia) share some physical traits, indicating
that the local H. erectus may have given rise to (or at least
contributed genes to) the modern humans in those regions. Although the
balance of the evidence at this point tends to favor the out of Africa
hypothesis, there is sufficient criticism of this evidence and the interpretation
of the evidence that a consensus has not yet been reached.
Cultural
Evolution
Culture
is the social transfer of information from one generation to the next.
Cultural evolution, or change in culture, is analogous to (but is not
the same as) biological evolution. Ideas change and these new ideas
are spread across populations. New ideas often compete with old ideas,
a form of natural selection. The evolution of language
had a huge impact on cultural evolution, speeding the rate of the spread
of ideas.
In a way, cultural evolution has “taken over” human populations.
Humans have not changed much genetically over the past 30,000 –
40,000 years, but culturally we are amazingly different from what we
were then. Culture can even influence biological evolution. For example,
when dairy cows were domesticated in some parts of the world, the people
in those populations eventually changed genetically. Adult humans in
most populations are unable to break down the sugar lactose, but in
populations where dairy cows were domesticated, the gene for lactose
tolerance was selected for and is now predominant. In other ways, culture
may be slowing down biological evolution. In our modern society, people
with genetic diseases that were probably selected against just a century
ago, due to death at a young age, are now able to live full lives and
even pass on those genes to their children. Culture dominates today’s
human society.
Useful
Websites
The
Institute of Human Origins
The
Smithsonian Institution Human Origins Program
PBS
Evolution
Lessons
On Human Evolution
There
are many nice classroom lessons about human evolution. The ENSI site
has many lessons, as well as a two-week unit that uses human biology
as an introduction to evolution, “Human
Evolution Unit.” This unit incorporates several of their
lessons on human evolution.
Skull
Comparisons
A
classic lesson on human evolution is the skull comparison, in which
students compare the skull characteristics of contemporary humans and
apes (such as chimps or gorillas), with fossil hominids. Students see
for themselves what kind of similarities and differences there are between
these species. There are a couple of lessons to choose from: ENSI’s
“Hominoid
Cranium Comparison,” as well as Access Excellence’s
“Hominoid
Skull Comparison.”
Chromosome
Comparisons
Another
common lesson is to compare the chromosome banding patterns of contemporary
humans and chimps (and sometimes gorillas). This is a nice visual way
to get across their extreme genetic similarity. ENSI has one: “Comparison
of Human and Chimpanzee Chromosomes.” PBS’
Evolution Teacher’s Guide has also adapted this lesson, called
“Chromosome
Clues.” Arizona State University’s Institute
for Human Origins also has an on-line activity, “The
Chromosome Connection: Comparison of Human and Ape Chromosomes.”
Phylogenies
There
are also several activities on phylogenies that include humans. ENSI’s
lesson,
“Classroom Cladogram of Vertebrate/Human Evolution,”
has the students collectively work together to build a giant phylogeny
of vertebrates, including humans. ENSI also has another activity, “Molecular
Sequences and Primate Evolution,” that has students
compare amino acid differences in beta hemoglobin from primate species,
and construct a phylogeny from these data. ASU’s Institute for
Human Origins has a lesson, “All
in the Family: Calculating Cousins” in which
students do not actually build a phylogeny, but they do examine evolutionary
relatedness, by calculating what they call “approximate cousinhood”
between humans and non-human primates. PBS’ Evolution web site
also has a web activity, “A
Tree Full of Ancestors,” in which students examine
three alternative phylogenies of extinct hominids, based on fossil data.
Real
Fossil Data
There
are several other lessons that use real fossil data to address questions.
PBS has a general lesson on hominid fossils, “Fossil
Finding,” in which students examine four real
fossil finds and learn how scientists interpret them and what kind of
questions can be answered by the fossils. ENSI’s “Chronology
Lab,” has students chart a timeline of fossil
hominids from fossil data. The teacher then makes a possible phylogeny
from these data, to help make the connection that these extinct species
are related. PBS’ Teacher’s Guide has a lesson, “Fossil
and Migration Patterns of Early Hominids” in
which students use real fossil data to determine possible migration
patterns of early hominids throughout the world.
Hominid
Adaptations
Finally, there are several lessons that explore important hominid adaptations.
Again, using real fossil data, there are two lessons that explore the
Laetoli footprints and bipedalism: ENSI’s “Footsteps
in Time,” in which students examine the fossil
prints, measure their own feet and height, then use the resulting correlation
to estimate the height of
the individuals who created the Laetoli print. PBS’
Teacher’s Guide’s “Watch
Your Step,” is a more general lesson on the prints,
exploring what kinds of questions can be addressed and how scientists
choose the best explanation for these kind of data. ASU’s Institute
for Human Origins has an on-line activity on bipedalism and the skeletal
structure that supports the ability to stand upright, “Building
Bodies: Primate Bipedalism—Understanding Standing Up.”
There is also a fun lesson on the key innovation of opposable thumbs,
“Thumbs
Up, Thumbs Down: Grasping the Idea of Evolution,”
in which students try various tasks first using their thumbs, and then
with their thumbs taped.
Videos
There
are also several videos you can use with your students. PBS’ Evolution
series, episode six, “The Mind’s Big Bang”
explores the later evolution of hominids, specifically the last 50,000
years or so of evolution in our own species. Also check out the PBS
video “Learning
and Teaching Evolution,” a companion to the series
that contains short, seven-minute segments for use in the classroom.
Segment five (“Did Humans Evolve?”) summarizes “The
Mind’s Big Bang.” Also check out Nova’s three-part
series: “In Search of Human Origins,” which
is a bit outdated as far as the most recent fossil evidence for our
earliest ancestors (it was filmed in 1994), but it has some nice segments
on the nature of this kind of historical science. ASU’s Institute
for Human Origins’ also has an online documentary, “Becoming
Human.”
Resource
List