Overview of Primate Evolution
Overview of Primate Evolution
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What is primate?
• Primates are difficult to define unambiguously in
terms of morphological attributes.
– Most primates have hands and feet adapted for
grasping.
– Relative to other mammals, they have large brains and
short jaws.
– They have flat nails on their digits, rather than narrow
claws.
– Primates also have relatively well-developed parental
care and relatively complex social behavior.
Primate Characteristics
• What is the major selective pressure associated with life in the trees?
• How are primates adapted to this life?
Binocular Vision
• Stereoscopic vision and resultant depth
perception allows primates to make accurate
judgments about distance and position of
adjoining tree limbs.
Mobile limbs
• Most primates have flat nails as well as sensitive pads
on the undersides of fingers and toes.
– Many also have both an opposable big toe and thumb
• Mobile limbs and clawless opposable digits allow
primates to freely grasp and release tree limbs
• Upright posture
• Small litter size
• Expanded forebrain, larger brain capacity
Primate origins
• The earliest primates were probably tree dwellers, shaped by natural
selection for arboreal life.
– The grasping hands and feet of primates are adaptations for hanging
on to tree branches.
• All modern primates, except Homo, have a big toe that is widely
separated from the other toes.
• The thumb is relatively mobile and separate from the fingers in all
primates, but a fully opposable thumb is found only in anthropoid
primates .
• Prosimians (~58mya)
• Monkeys (35mya)
• Apes (23mya)
• Hominids (~6mya)
First Arboreal Mammals
Ring-tailed lemur
Potto • The oldest known anthropoid fossils, from
about 45 million years ago, support the
hypothesis that tarsiers are the prosimians
most closely related to anthropoids
Prosimians: Lorises, Lemurs
lorises
10 cm
Lemurs
Tarsier
ANTHROPOIDS
2. Anthropoids = monkeys + apes + man
• Monkeys By about 40 million years ago, – Old world
(Cercopithecoids)
monkeys include arboreal
– All New World monkeys were established in Africa, Asia, (Old and ground-dwelling
monkeys
(Ceboids) are World) and South America (New World) species
arboreal – Old world species lack
– New World
• Old world monkeys prehensile tails and their
monkeys have nostrils open downward.
prehensile tails • New world monkeys
and nostrils that
open to the side
• With the exception of gibbons, modern apes are larger than monkeys, with relatively long
arms and short legs, and no tails,
• Although all apes are capable of brachiating, only gibbons and orangutans are primarily
arboreal.
• Social organization varies among the genera, with gorilla and chimpanzees highly social.
• Apes have relatively larger brains than monkeys, and their behavior is more flexible.
Skull Comparisons:
Timing divergence from the molecular clock and fossils
Human Origins
• In the continuity of life spanning over 3.5 billion years, humans and apes
have shared ancestry for all but the last few million years.
• two words that are easy to confuse but which have distinct meanings.
– Hominoid is a term referring to great apes and humans collectively
– Hominid has a narrow meaning, confined to those twigs of the
evolutionary tree that are more closely related to us than any other living
species.
• There are two main groups of hominids: the australopithecines, which
came first and are all extinct, and members of the genus Homo, with all
species extinct except one: Homo sapiens.
Misconceptions about human
origins• First, our ancestors were not chimpanzees or any other modern apes.
– Chimpanzees and humans represent two divergent branches of the hominoid
tree that evolved from a common ancestor that was neither a chimpanzee nor a
human.
• Second, human evolution did not occur as a ladder with a series of steps leading
directly from an ancestral hominoid to Homo sapiens.
– If human evolution is a parade, then many splinter groups traveled down dead
ends and several different human species coexisted.
– Human phylogeny is more like a multibranched bush with our species as the tip
of the only surviving twig.
• Third, the various human characteristics, such as upright posture and an enlarged
brain, did not evolve in unison.
– Different features evolved at different rates, called mosaic evolution.
To be Human
• standard primate characteristics
• upright, bipedal posture
• much expanded brain
• well defined sexual dimorphism
• extended childhood
• language
• society
• culture
Primate Characteristics/Trends
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Human Ancestors: Hominids
6-7 mya
2002 discovery of hominid from Chad with a
mosaic of primitive (very small brain) and derived
(small canines) features.
Earliest hominins: pre-Australopiths
•Sahelanthropus tchadensis. (Toumai, “hope of life” in Goran). A single
skull, jaw fragments, several teeth, unearthed in 2002 by Michael Brunet,
dated to about 6.5 mybp Found in Chad, central Africa?
•Forward position of foramen magnum suggest bipedalism
▪ Orrorin tugenensis Femoral
Morphology and the
Evolution of Hominin
Bipedalism
Bipedalism is a key human adaptation and a defining feature of the hominin clade. Fossil femora discovered in Kenya and
attributed to Orrorin tugenensis, at 6 million years ago, purportedly provide the earliest postcranial evidence of hominin
bipedalism, but their functional and phylogenetic affinities are controversial. We show that the O. tugenensis femur differs
from those of apes and Homo and most strongly resembles those of Australopithecus and Paranthropus, indicating that
O. tugenensis was bipedal but is not more closely related to Homo than to Australopithecus. Femoral morphology
indicates that O. tugenensis shared distinctive hip biomechanics with australopiths, suggesting that this complex evolved
early in human evolution and persisted for almost 4 million years until modifications of the hip appeared in the late
Pliocene in early Homo.
Science 21 March 2008:
Vol. 319. no. 5870, pp. 1662 - 1665
Earliest hominins: pre-Australopiths
Ardipithecus ramidus
▪ 4.5 mya
• Robust ones have large premolar & molar teeth and sagittal crest
Hominid Fossil Record
• All known hominid (human) fossils older than about 1.5 million years are from
eastern and southern Africa.
– Most consist of teeth and fragments of jaws, skulls, and other skeletal pieces,
with a few spectacular exceptions.
– Researchers try to reconstruct human phylogeny from an incomplete record,
revising their hypotheses to account for new fossil evidence and data from new
research strategies such as molecular systematics (DNA and Protein studies).
• The various pre-Homo hominids are classified in the genus Australopithecus
(“southern ape”) and are known as australopithecines.
– The first australopithecine, A. africanus, was discovered in 1924 by
Raymond Dart in a quarry in South Africa.
• From this and other skeletons, A. africanus probably walked fully erect
and had human-like hands and teeth.
• However, the brain was only about one-third the size of a modern
human’s brain.
• In 1974, a new fossil, about 40% complete, was discovered in the Afar region of
Ethiopia.
– This fossil, nicknamed “Lucy,” was described as a new species, A.
afarensis. It was discovered by Don Johanson and his team of
paleoanthropolgists.
Hominid Fossil Record
• In the past few years, paleoanthropologists have found hominid species that
predate A. afarensis.
– The oldest fossil that is unambiguously more human than ape is
Australopithecus anamensis, which lived over 4 million years ago.
– Other fossils of putative hominids go back 6 million years, closer to the ape-
human split that molecular systematists estimate occurred about 5 - 7 million
years ago.
• The australopithicines were evolutionary dead ends. The question remains which
were either on, or close to, the phylogenetic lineage that led to the Homo branch.
– Two lineages appeared after A. afarensis: the “robust” australopithecines
with sturdy skulls and powerful jaws and teeth for grinding and chewing
hard, tough foods (A. boisei “Nutcracker man”); and the “gracile”
australopithecines with lighter feeding equipment adapted for softer
foods.
– Most researchers agree that the robust australopithecines were an
evolutionary dead end, and that the ancestors of Homo were among the
gracile australopithecines.
Map of Australopithecine Finds
Map of Australopithecus
sites in Africa, with a
focus on the East African
rift valley and limestone
caves of South Africa.
Hominid Fossil Record: “Lucy”
Lucy
• Based on this fossil and other
discoveries, this species had a
brain the size of a chimpanzee, a
prognathic jaw, longer arms (for
some level of arboreal
locomotion), and sexual
dimorphism more apelike than
human. Modern Human Chimp
– However, the pelvis and skul
bones and fossil tracks showed
that A. afarensis • Lack of a splayed big
toe suggests full
walked bipedally. bipedalism by 3 mya
o Discovered in Ethiopia in 1974.
o Dated at 3.2 million years old!
o 40% of her skeleton was found.
o Only four feet tall.
o Bipedal for certain. She walked
upright.
An early Australopithecus afarensis postcranium from
Woranso-Mille, Ethiopia
Haile-Selassie et al.
▪ Small brains
• Homo rudolfensis: 1970’s Richard Leakey. Brain size 750cc, but with
more primitive looking face. Both dated to around 2.3-2.0 mybp
Fossil Record of Genus Homo
• The earliest fossils that anthropologists place in our
genus, Homo, are classified as Homo habilis. The
“Tool making man”
– These fossils range in age from 2.5 to 1.6 million
years old.
– This species had less prognathic jaws and larger
brains (about 600 - 750 cm3) than
australopithecines.
– In some cases, anthropologists have found sharp
stone tools with these fossils, indicating that
some hominids had started to use their brains
and hands to fashion tools.
• A remarkably complete fossil of a young hominid
known as “Turkana Boy” indicates that even larger
brains had evolved by 1.6 million years ago.
– The body had a brain that
would probably be over
900 cm3 in an adult of his
species, a size between Turkana Boy
that of H. habilis and
H. erectus.
Fossil Record: Genus Homo
• Homo erectus was the first hominid
species to migrate out of Africa,
colonizing Asia and Europe.
– They lived from about 1.8 million
to 500,000 years ago.
• Fossils from Asia are known
by such names as “Peking
man” and “Java Man”.
• In Europe, H. erectus gave rise
to the humans known as
Neanderthals.
– Compared to H. habilis, H.
erectus was taller, had a larger
brain (averaging about 1,100
cm3), and had about the same
level of sexual dimorphism as
modern humans.
– They were most successful of all
species of the genus Homo
because they existed as a
successful species for more than
1 million years!
Homo erectus/ergaster
• Larger tools, more
specialized, exemplified by
the hand axe
• Later versions of hand axe
(5mybp) suggest an image to
guide tool creation, attention
to symmetry and shape.
Some may not have been
strictly utilitarian.
• African ergaster:
– Ergaster migrated out of African almost as soon as it emerged.
– The ergaster that remained in Africa either (a) slowly evolved into
modern Homo sapiens or (b) went extinct, leaving Africa to be re-
colonized by descendants of Asian erectus who eventually evolved
into modern Homo sapiens
A tale of two species: African ergaster and Asian erectus
• Asian erectus
remained in extreme southeast Asia until as recently as 20,000 ybp
(Homo florensiensis – ‘the hobbit’)
Evolved into Homo heidelbergensis in Eurasia (500,000ybp) who
eventually gave rise to Neanderthals and quite possibly Homo sapiens
H. heidelbergensis very likely migrated extensively throughout
Europe, east Asia, and possibly Africa. Probably the first true big-
game hunter.
Migrate Out-of-Africa
• The earliest specimens of Homo erectus are found in Africa, but,
sometime after 1 million years ago, Homo erectus apparently migrated out
of Arica.
• The earliest Homo erectus finds are in the Rift Valley of Africa and in South
Africa.
• Stone tools and camp sites are widely distributed over Africa, including
sites in what is now the Sahara desert. By at least 1 million years ago, H.
erectus migrated out of Africa to Asia and Europe.
• Tools and remains of this species have been found widely distributed in
Europe and Asia.
• Homo erectus is thus the first human species to migrate out of Africa and
adapt to a variety of Old World environments.
• Recent dating techniques applied to earlier finds in Java and the Caucasus
mountains indicate dates much earlier than that and would place Homo
erectus in Asia over 2 million years ago.
– Such early dates would, if proven accurate, dramatically overturn the
currently accepted chronology of early human development.
– At this date, however, the earliest claimed dates for H. erectus outside
of Africa are still highly controversial.
Homo ergaster (Narikatome Boy)
• By 1.9 million years ago, another lineage of the genus Homo emerged in Africa. This species
was Homo ergaster.
• Traditionally, scientists have referred to this species as Homo erectus and linked this species
name with a proliferation of populations across Africa, Europe, and Asia.
• Yet, since the first discoveries of Homo erectus, it had been noted that there were differences
between the early populations of "Homo erectus" in Africa, and the later populations of Europe,
Africa and Asia.
• Many researchers now separate the two into distinct species Homo ergaster for early African
"Homo erectus", and Homo erectus for later populations mainly in Asia.
• Since modern humans share the same differences as H. ergaster with the Asian H. erectus,
scientist consider H. ergaster as the probable ancestor of later Homo populations.
• Boy, 12 years
• Long legs narrow hips and shoulders
• Vertebrate opening in spinal cord smaller than moderns
• Brain size large (850-1100)
• Front tooth size increase, back tooth decrease
• Tall (5-5.5 feet), thick bones.
What happened?
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Human Phylogeny
LUCY
Human Evolution
Summary of fossil evidence on the recent ancestry of humans
Origin of Homo sapiens
• Two alternative hypotheses have been
proposed for the origin of anatomically
modern humans.
• In the multiregional
hypothesis, fully modern
humans evolved in parallel
from the local populations
of H. erectus.
– The multiregional hypothesis places the last
common ancestor in Africa over 1.5 million
years ago, when H. erectus began migrating
to other parts of the world about 100,000
years ago.
• A compromise alternative to these extremes suggests that Homo sapiens originated and then dispersed from Africa
100,000 years ago.
– These individuals then interbred with the regional descendents of the earlier H. erectus migration.
– This hypothesis predicts that the genomes of indigenous people from around the world today should reflect a
complex mix of ancestries.
1. Which Hominid species gave rise
to Homo sapiens?
Homo sapiens Out of Africa
• So far, the genetic data have mostly supported the replacement hypothesis.
– Using changes in mitochondrial DNA (mtDNA) among human populations as a molecular clock,
research have reported a time of genetic divergence of about 100,000 years ago.
• This is supported by nuclear genetic markers.
– The mtDNA extracted from Neanderthal bones fall completely outside the range of mtDNA for
modern Europeans.
• These data suggest that Neanderthals contributed nothing to the ancestry of anatomically modern humans
in Europe.
• So far, the fossil evidence has been less one-sided than the genetic data in testing
the alternative hypotheses.
– The western European fossil evidence is consistent with total replacement of Neanderthals about
40,000 years ago by anatomically modern humans, known as Cro-Magnons.
• There were no intermediates suggesting interbreeding between Neanderthals and the later arrivals.
– However, fossil evidence from outside Europe is more ambiguous, with some paleoanthropologists
interpreting some Asian fossils as intermediates between older fossils of H. erectus and the
skeletal features of modern Asians.
Why was Homo erectus so successful?
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Trends in
Skull
Development
Cro-Magnon Man
Cro-Magnon humans
35,000 years B.P. in western Europe to 17,000
years B.P.
1,600 cc cranial capacity
Name comes from a hotel in France
Not a different species, just old Homo sapiens
from Europe
Shelters
• 15,000 y BP Ukraine
• Some made with mammoth bones
Wood, leather working; carpentry
• Tools
• From cores to blades
Specialization
• Composite tools
• Bow and arrow
Domestication of dogs
• Gathering rather than hunting
Cro-Magnon Man – Homo sapiens
Cave paintings
Mostly animals on bare walls
Subjects were animals favored for their meat
and skins
Human figures were rarely drawn due to
taboos and fears that it would somehow
harm others
Language and Cultural Evolution
Up the ladder
The idea developed that more than
one hominid species existed at the
same time developed
A. robustus
Homo habilis
Homo erectus
Australopithecus
africanus
Australopithecus
Australopithecus afarensis Added
“Lucy” 1974
Branching Out
1991 Changing status
Homo sapiens
Homo erectus
As more specimens
were found a clearer A. robustus A. africanus
idea developed of the
relationships between
them
Australopithecus afarensis
2001 From a tree to a bush
Homo sapiens
Homo neanderthalensis
Homo
H. heidelbergensis erectus
1 Ma
Australopithecus ?
4 Ma afarensis
Australopithecus anemensis
? ?
Ardipithecus ramidus
2003 Gorilla gorilla Pan trogolodites
Homo neanderthalensis
Homo sapiens
DEEPER
Homo
ROOTS H. heidelbergensis erectus
1 Ma
H. ergaster
Paranthropus
Paranthropus. robustus
boisei ? Homo
2 Ma Homo habilis
rudolfensis
A. africanus
P. aethiopicus
A. garhi
?
3 Ma
Australopithecus
afarensis ?
4 Ma Australopithecus anemensis
?
? Ardipithecus
5 Ma
ramidus
?
6 Ma Orrorin tugensis
?
?
Sahelanthropus tchadensis
“Toumai”
7 Ma
Human evolution
3) 200 kyr:
Genus Homo sapiens
Modern man
2) 2 Myr
Genus Homo erectus
Stone tools
1) 4-6 Myr:
Genus Australopithecus
Walk upright
DNA
Out of Africa Dispersals
Molecular studies suggests that Africans exhibit a higher level of genetic diversity than non-Africans
for
many mitochondrial and nuclear loci. This implies that the African line has been around for longer
than other
populations, enabling greater genetic variation to accumulate,which is in agreement with the OAR
mode
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Adaptive Changes in Pliocene Hominins
• Hominins may have begun in the Miocene age, before the Pliocene,
several hominin lineages originated and went extinct during the epoch
although exact dates of everything is very uncertain.
Examples of hominins during era: Homo, Australopithecus and
Paranthropus
1. Mobility: known due to changing limb proportions, object transport
by toolmakers and biogeographic spread between regions.
2. Foraging: known by accumulation of flaked stone and hammers.
3. Diet: known by changing dental proportions, use of tools to access
food from large animals.
Habitat of Hominins
• Different Hominins species lived quite differently, early in the Pliocene period
hominins lived more in woodland and forest, later on a shift towards open
vegetation.
• This change in habitat which effected their evolution was linked to climate.
• There are specific adaptations that can be associated with changes in environment
even though it is hard to go from correlation to cause effect.
• Certain characteristics seen in Hominins are thought to be due to “Variability in
environment
Variable Selection: adaptation evolved due to increased environmental variability.