Chapter 20– Human Evolution

“Light will be thrown on the origin of man and

his history” -Darwin 1859, p. 488

I. Relationships among Humans and the Extant

Apes
II. The Recent Ancestry of Humans
III. The Origin of the Species Homo sapiens
IV. The Evolution of Distinctive Human Traits (on
your own)
• Order Primates = lemurs, monkeys, apes
• Suborder Catarrhini = Old World monkeys & apes
• Superfamily Hominoidea = great and lesser apes
• Family Hominidae = great apes and humans
• Hominins = anything more closely related
to Homo sapiens, the anatomically modern
human, than to other apes
I. Relationships among Humans and the Extant Apes
Humans are in the same clade as the African great Apes
• Synapomorphic characters with the Apes– large brain,
absence of tail, erect posture, flexible hips and ankles,
wrist, and thumb, structure of arm/shoulder
• More synapomorphies are
shared with the African Great
Apes– elongated skull, enlarged
brow, shortened canine teeth,
upper-jaw changes, fusion of
wrist bones, enlarged ovaries
and mammary glands, muscle
anatomy, and reduced hairiness
 • Molecular analyses of human and primate blood
serum albumins also shows that humans are most
closely related to the African Great Apes

• These data suggest a divergence date of ~5mya– but the

polytomy among humans, gorillas, and chimps remains
Humans, Gorillas, Chimpanzees and Bonobos

• The four ways

the polytomy
could be
resolved

The consensus among scientists is that the most likely

tree topology is that shown in (a)– where humans are
most closely related to chimpanzees and bonobos
Molecular Evidence
• Using the DNA sequence of a mitochondrial gene
reveals that humans are most closely related to the
common chimpanzees

• Most molecular
analyses show
similar
relationships
among humans
and the great
apes, but not all
• Some molecular analyses have produced phylogenies
where chimps and gorillas, or gorillas are the closest
living relatives to humans

• Remember, gene trees do not always reflect the species

tree—incomplete lineage sorting
• Entire genome analysis shows that for most genomic
regions humans and chimps are most closely related
Morphological Evidence
• Several features shared by chimps. and gorillas are
missing in humans–
• Knuckle walking skeletal traits
• Either this characteristic was derived once, prior to
the gorilla/chimp/human split, and was lost in the
human lineage OR…
• The human lineage diverged from the gorillas and
chimps first, and the trait was derived only in the
lineage leading to the chimps and gorillas

• Although the second possibility is the most

parismonious for this trait, it would complicate the
parsimony of many other morphological characters
• Difficult to identify which characters are ancestral or
derived
• Dryopithecus is an extinct European ape 10 million
years old
• Shares many traits with gorillas that humans and
chimps lack

• If these characters are ancestral and humans and

chimps don’t have them, it is most parsimonious for
humans and chimps to be sister species
• Estimation of the divergence times of the great ape
species– based upon molecular clock estimates from
dozens of genes

• Gorillas diverged ~6.4 mya

• Humans and chimps split ~5.4 mya
Genetic Differences between Humans, Chimpanzees, and
Gorillas

• Chimps and gorillas have 24 pairs of chromosomes

• Human ch. 2 is a fusion of 2 ancestral chromosomes
• Genetic losses and additions in humans
• Loss of androgen receptor that
produces whiskers
• Addition of an allele via transposable
element relocation—brain-expressed
protein
Examination of whole genome organization shows how
little difference there is between humans and chimps
• Many biologists argue that both should be in the
genus Homo

• ~80 genes are active in

chimps but not in humans
• Olfactory, taste,
immune response
• Human specific gene
SRGAP2C—extends brain
development and
increases dendrite
development
Many other differences lie in gene regulatory regions and
processes
• miR-320b is a micro RNA

• Humans have significantly

increased expression of the
miRNA and therefore
significantly less expression of
the 18 targets of miR-320b in
cells of the prefrontal cortex
II. The Recent Ancestry of Humans

• Most recent common ancestor

likely was:
• A knuckle walker, had a broad
fruit-based diet, lived in a wide
range of habitats, probably
used tools, hunted, and likely
had culture

• Is the history of human evolution a

single lineage of slow
transformation, or a radiant one
with extinctions and splits?
• Fossil Evidence and Interpretation
• The fossil record includes a diversity of hominins—
species that lived after the human and chimpanzee
lineages separated and are more closely related to
humans than to chimpanzees
• Sahelanthropus tchadensis
• At 6-7 million years old, this is the earliest known
record of the human family
• Discovered in Chad in Central Africa, this skull,
nicknamed 'Toumaï', comes from a crucial yet little-
known interval when the human lineage was
becoming distinct from that of chimpanzees
• The skull has a combination of primitive and
advanced features
• Small brain case—looks chimp from behind
• Flatter face that’s more Homo-like
• Perhaps the “missing link” between hominins and
other apes—could be the last common ancestor
between humans and the chimpanzees
 Australopithecines are among the earliest humans

• Two main body forms

• Robust and gracile
• Bipedal like modern humans
• Known from limb structure
and footprints
• Australopithecines were short with small brain cases

• Modern human
structure more similar
to gracile
australopithecines
• Australopithecus

• Expect that they lived

until less than 2mya
Paranthropus
• Formerly known as ‘robust australopithecines’

• had enormous teeth

and very strong jaw
muscles for biting tree
branches (like gorillas
do today)
• Bipedal
• First Homo was H. habilis
• Homo had larger brain cases and smaller teeth than
Australopithecus
• Homo was also taller with less sexual dimorphism in
size
• Homo rudolfensis may be the same species as H.
habilis but have some skeletal differences

• H. habilis have
been found with
stone tools and
animal bones
indicating butchery
• H. ergaster is a more
recent ancestor from
Africa
• H. erectus was first
human to leave Africa
• H. heidelbergenesis is its
descendent
• H. neanderthalensis may
be our ancestor or may
have been a sister species
that died out

• All have smaller faces and much larger brain cases than
previous hominins
• Anatomically modern Homo sapiens appeared
approximately 100,000 ya
• Prior to Homo sapiens, as many as 5 human species
existed at the time
• Some in the same regions
• How do we determine the relatedness of human
species?
Summary of fossil evidence on the recent ancestry of
humans
• Cladogram of the • Phylogeny of Homo
extinct homonins sapiens and recent
and modern apes ancestry
III. The Origin of the Species Homo sapiens

• Paleontologists disagree about the

taxonomic status of H. ergaster
• Same species as H. erectus or different?
• New H. erectus fossil recently found in
Georgia from 1.75 Mya
• There is also disagreement about status
of H. neanderthalensis
• Species or subspecies of H. sapiens?
 • General agreement that H. sapiens are descendant
from H. ergaster/erectus
• All homonins prior to ergaster/erectus were confined
to Africa

• Three
hypotheses
as to how
ergaster/
erectus
evolution
occurred
• African replacement model– H. sapiens evolved in
Africa and subsequently radiated to Asia and Europe
• Replacing erectus and neanderthalensis without
interbreeding

• Multiregional evolution model--H. erectus migrated to

and the slowly diverging populations continued to
interbreed
• Hybridization and assimilation model—H. sapiens
evolved in Africa and migrated to Asia and Europe but
also interbred with H. neanderthalenis, and H. erectus
African Replacement versus Multiregional Evolution:
Archaeological and Paleontological Evidence
• African replacement model contends that long
established populations of H. erectus/H. neanderthal
were ‘replaced’ throughout Europe and Asia by H.
sapiens without interbreeding

• Would have required direct competition between

the two for resources

• Much fossil evidence supports the mutli-regional

evolution– at least with respect to Europe
• Molecular Evidence
• Phylogenetic and genetic expectations of the two
models
• Comparison of complete mtDNA from 53 humans

• All non-African
individuals branch
from within the
Africans

• Supports the African

replacement model
for mitochondrial
genes
•
• However, gene flow between populations
could cloud this conclusion

• Species can’t diverge before alleles diverge,

unless there is continued gene flow
• Analysis of multiple loci--30 microsatellite
loci from 14 populations of humans
• Again- support of
the African-
replacement
model
• Analysis of genetic diversity among
humans from 7 different geographical
regions
• Diversity like that seen in founding
events– the founder effect
• Only a portion of the original genetic
variation is carried to each new location

• Consistent with the African replacement

model

• Estimated that modern humans left Africa

about 100,000 years ago
 African Replacement versus Multiregional Evolution:
Molecular Evidence from Premodern Humans
• Paabo recovered genome sequences of three H.
neanderthalensis bones
• Also a Denisovan specimen
• An undescribed human species from Siberia about
30,000–40,000 ya
• Found that modern
humans are more
closely related to
each other than
any is to H.
neanderthalensis
or the Denisovan
fossil
Green did a SNP study with the same data that could
detect gene flow
• Found that modern humans contained SNPs from
neanderthals and Denisovans
• Supports Hybridization and Assimilation model
• “Leaky replacement”
 Genetic Diversity among Living Humans

• All currently living humans

have extreme genetic
similarity and most
variation can be found
within populations rather
than between populations
• Using massive data sets, we can distinguish
geographical groupings of humans