Human evolution 1 Classification of Hominins Kingdom: Animalia Subphylum: Vertebrata Class: Mammalia Order: Primates Suborder: Anthropoidea Superfamily: Hominoidea Family: Hominidea Subfamily: Homininae Tribe: Hominini 2 T. Harrison Science 327, 532-534 (2010) Animalia: depend on intake of living food; do not make their own food Subphylum: have a bony skeleton with bilateral symmetry Class: live bearing, mother nurses young, 4 chambered heart, warm blooded, sweat glands, 2 sets of teeth with four kinds of teeth Order: Generalized arboreal; 5 digits, hands and feet capable of grasping; tendency to erect posture with head balanced on spinal column; reliance on vision rather than smell; 3D vision; nails rather than claws; opposable thumb; extensive bony protection of eyes Suborder: New World monkeys, Old World monkeys, apes, and humans; are two other primate suborders (prosimians and tarsiers) Superfamily: short, flat, broad trunk; vertebral column set within rib cage; dorsal scapula & lateral facing shoulder = full rotation of arms; no external tail; specialized dentition Family: humans, ancestors, & all great apes Subfamily: humans, ancestors, chimps, & gorillas Tribe: humans and ancestors 2 What is a Hominin? Modern humans Our ancestors Other species branching off this lineage First Hominin debate 5 – 10 mya Earliest Hominin fossils are: African, ~7 million years old Very apelike – mosaic of traits 3 At the end of the Miocene, around 6 mya, a new tribe of primates arose: the Hominins. This family includes modern humans and our extinct ancestors since the divergence from the last common ancestor with chimpanzees. DNA evidence and protein studies suggest that the last common ancestor to the chimpanzee and human lineages evolved at this time, around 6 mya; however, this evidence does not tell us about what these early ancestors would have looked like  Hominins, members of the taxonomic family Homininae, may be distinguished from the apes by the following anatomical traits: (next slide) 3 Identifying Hominins Key Hominin traits: Skeletal traits associated with bipedalism Changes to the face, teeth, and jaws Expansion of the brain (encephalization) Tool-making behavior 4 The identification of hominin fossils depends on some key skeletal traits BUT identifications are not certain in all circumstances. (Why is this the case? Well recall our discussion in class regarding the fossil record for primates…). The key traits we look at are: - bipedalism -dental traits -increased cephalisation, or increased brain size -a reduced face, also known as a reduction in facial prognathism -and finally a little bit of culture It is important to realize that many of these traits evolved in a mosaic fashion. Many of the early hominids that we will be discussing today will not show all of these traits but a subset of them. This can cause controversy in classification, as different researchers emphasize th ...

1. Human evolution
1
Classification of Hominins
Kingdom: Animalia
Subphylum: Vertebrata
Class: Mammalia
Order: Primates
Suborder: Anthropoidea
Superfamily: Hominoidea
Family: Hominidea
Subfamily: Homininae
Tribe: Hominini
2
T. Harrison Science 327, 532-534 (2010)

2. Animalia: depend on intake of living food; do not make their
own food
Subphylum: have a bony skeleton with bilateral symmetry
Class: live bearing, mother nurses young, 4 chambered heart,
warm blooded, sweat glands, 2 sets of teeth with four kinds of
teeth
Order: Generalized arboreal; 5 digits, hands and feet capable of
grasping; tendency to erect posture with head balanced on
spinal column; reliance on vision rather than smell; 3D vision;
nails rather than claws; opposable thumb; extensive bony
protection of eyes
Suborder: New World monkeys, Old World monkeys, apes, and
humans; are two other primate suborders (prosimians and
tarsiers)
Superfamily: short, flat, broad trunk; vertebral column set
within rib cage; dorsal scapula & lateral facing shoulder = full
rotation of arms; no external tail; specialized dentition
Family: humans, ancestors, & all great apes
Subfamily: humans, ancestors, chimps, & gorillas
Tribe: humans and ancestors
2
What is a Hominin?
Modern humans
Our ancestors
Other species branching off this lineage
First Hominin debate
5 – 10 mya
Earliest Hominin fossils are:
African, ~7 million years old
Very apelike – mosaic of traits

3. 3
At the end of the Miocene, around 6 mya, a new tribe of
primates arose: the Hominins.
This family includes modern humans and our extinct ancestors
since the divergence from the last common ancestor with
chimpanzees.
DNA evidence and protein studies suggest that the last common
ancestor to the chimpanzee and human lineages evolved at this
time, around 6 mya; however, this evidence does not tell us
about what these early ancestors would have looked like
Hominins, members of the taxonomic family Homininae, may
be distinguished from the apes by the following anatomical
traits: (next slide)
3
Identifying Hominins
Key Hominin traits:
Skeletal traits associated with bipedalism
Changes to the face, teeth, and jaws
Expansion of the brain (encephalization)
Tool-making behavior
4

4. The identification of hominin fossils depends on some key
skeletal traits BUT identifications are not certain in all
circumstances. (Why is this the case? Well recall our discussion
in class regarding the fossil record for primates…). The key
traits we look at are:
- bipedalism
-dental traits
-increased cephalisation, or increased brain size
-a reduced face, also known as a reduction in facial prognathism
-and finally a little bit of culture
It is important to realize that many of these traits evolved in a
mosaic fashion. Many of the early hominids that we will be
discussing today will not show all of these traits but a subset of
them. This can cause controversy in classification, as different
researchers emphasize the importance of certain traits over
others.
So these characteristics are what makes a hominin a hominin;
however, it is important to recognize that not all these
characteristics developed simultaneously or at the same pace. In
the earliest stages of hominid evolution, skeletal evidence
indicating bipedal locomotion is the only truly reliable indicator
that fossils were hominids. Later, other features, as those
mentioned here, become highly significant.
4
First Some Review:
How Do Primates Get Around?
Primates exhibit several locomotory patterns:
Vertical Clinging and Leaping
Brachiation

5. Quadrupedalism
Arboreal vs. terrestrial
Knuckle walking
Bipedalism
Habitual
5
-most primates use more than one form of locomotion and they
owe this important ability to their generalized anatomy
-I’m just going to go over the 3 most common forms of
locomotion, but of course, other forms do exist
-before I go into this you must remember that it is difficult to
lump a primate species into one group, since I’ve already said
that most primates practice a combination of several modes of
locomotion
-we will be going through this in lab and looking at the
skeletons of several primates to see their locomotory adaptation,
so I’m going to go over it quickly
-so, now, we will be looking at vertical clinging and leaping,
brachiation, arboreal and terrestrial quadrupedalism
-and because it’s how we get around and seems to be a key way
we are different from all other living primates, we’ll cover
bipedalism this topic
5
How Do Nonhuman Primates Get Around?

6. Vertical Clinging and Leaping
Used by many lemurs, lorises, galagos (strepsirrhines) and the
tarsiers
Grasp onto trunks
Spring off using their long hind limbs
Hindlimbs > Forelimbs
6
-this form of locomotion is characteristic of many lemur,
lorises, and tarsiers
-as the term implies, vertical clingers and leapers support
themselves vertically by grasping onto trunks of trees or other
large plants while their knees and ankles are tightly fixed
-by forcefully extending their long hind limbs, they can spring
powerfully away either forwards or backwards
-we will be examining the bones of several vertical clinger and
leapers,
-you will notice that their hindlimbs are longer than their
forlimbs to allow the massive leap
6
How Do Nonhuman Primates Get Around?
Brachiation
Also known as “arm swinging”
A suspensory form of locomotion
Long curved fingers & other modifications to shoulder
Forelimbs > Hindlimbs
e.g., gibbons

7. 7
-brachiation, or arm swinging, is a suspensory form of
locomotion in which the body moves by being alternatively
supported under either forelimb
-you may have brachiated as a child on “monkey bars” in
playgrounds
-because of anatomical modifications at the shoulder joint, apes,
and humans are capable of true brachiation
-however, only the small gibbons and siamangs of Southeast
Asia use this form of locomotion almost exclusively
-brachiation is seen in species characterized by arms longer than
legs and long curved fingers
7
How Do Nonhuman Primates Get Around?
Quadrupedalism
Using all four limbs to support the body during locomotion
Arboreal vs. terrestrial
Knuckle walking (Figure 6.28)
Modifications for each form
Forelimbs ~= Hindlimbs
8

8. -almost all primates are, at least to some degree, quadrupedal,
meaning they use all four limbs to support the body during
locomotion
-most of the quadrupedal primates are primarily arboreal but
terrestrial quadrupedalism is also common
-the limbs of terrestrial quadrupeds are approximately the same
length, with forelimbs being 90% or more as long as hind limbs
-in arboreal quadrupeds, forelimbs are somewhat shorter
-knuckle walking is a sub-category of quadrupedalism found in
chimpanzees, bonobos and gorillas.
-These animals walk on the knuckles of their hands and the
soles of their feet
-the forelimbs are slightly larger than the hindlimbs
8
The Bipedal Adaptation
We, hominins, are unique!
We are fully bipedal, and not good at other forms of locomotion
We are pretty specialized for bipedalism
What are some of the skeletal changes that took place to allow
for bipedalism?

9. 9
-we are unique among primates because we are fully bipedal and
we’re not really good at other forms of locomotion
-so, we are definitely using a very specialized form of
bipedalism
-compared to other primates that only walk bipedally every now
and then, or occasionally to perform specific tasks
-and when they do it, it’s pretty inefficient
-our mode of locomotion, bipedalism, is pretty extraordinary
when you think about it
-the act of human walking is almost like falling repeatedly
-the problem is to maintain balance on the stance leg while the
swing leg is off the ground
-so, for us to become bipedal, it required many changes from
the typical monkey/ape pattern
-so, what are some of these changes?
9
The Skeletal Evidence of Bipedalism
https://youtu.be/Y_o9QFWSTbA

10. Flaring of hips for weight distribution
S-shaped curve to spinal column
Double arched foot
Change in limb proportions
Modifications also occurred in other parts of the skeleton
because of the shift to bipedalism. The most significant of these
include:
- The repositioning of the foramen magnum (the foramen
magnum is the hole at the base of our skull where our spinal
cord enters our skull to connect with our brain). In hominins it
is moved forward so it is directly underneath the skull and the
head is more or less balanced on the spine. In quadrupeds it is
towards the back of the skull.
- The addition of spinal curves that help to transmit the weight
of the upper body to the hips in an upright position
- The shortening and widening of the pelvis (this assists with
supporting our massive body weight on top of our two lower
limbs)
- The lengthening of the hind limbs, which increased stride
length
- The angling of the femur inward to bring the knees and feet
closer together under the body
- There are also several structural changes in the foot, including
the development of the longitudinal arch and the realignment of
the big toe in parallel with the other toes; the longitudinal arch

11. helps to absorb shock and adds a propulsive spring
10
Why Bipedalism ?
Freeing of the hands
Prolonged infant care
Food carrying or provisioning activities
Tool use
Elevate head
Visibility in open grasslands
Energetic advantages
Long distance, slow walking
Improved heat regulation
Mating advantage?
Lower body stresses
Knee, hip, and lower back pain
Advantages
Disadvantages (Costs)
11
Why become bipedal? This is a key question, because
bipedalism is what first set our lineage apart from other
primates. What selection pressures favored bipedalism? there
are many ideas, no consensus, and many of the ideas are flawed;
of course, there may have been multiple reasons why better
bipeds left more offspring
Most of the theories have to do with these three ideas:

12. -Energy efficiency: researchers believe that bipedal walking is
just more efficient in terms of energy expenditure in comparison
to quadrupedalism; so, from knuckle walking to bipedalism
would have been a good jump
-Vertical posture exposes less of the body to direct sun.
Bipedalism holds the body vertical, instead of horizontal as in
quadrupedalism, so the sun strikes a smaller fraction of the
body when it is upright. This means bipedalism may have been
favoured because it helped animals in increasingly open
grasslands to avoid overheating (and increased distance from
the ground facilitates cooling by increased exposure to breezes)
-However, we don’t know if the adaptation to bipedalism
happened on the grasslands, could have been in the woodlands
where there is shade
-Upright posture provided access to seeds, berries, etc. in higher
branches in trees and standing up provided a better view of the
surrounding countryside; this would give them a view of
potential predators as well as other group members
-Upright posture free the arms to carry various objects (but
chimps carry things in their hands reasonably well even without
being good bipeds)
-Others have suggested that bipedalism offers mating benefits
as it is easier to make a social display if you can walk upright;
the more upright you are the more imposing you look, could
lead to more mating success
Obviously, all of these theories have flaws
11
Dating Techniques

13. For a longer instructional video see:
https://youtu.be/gqDHzCj90co
When Did the First Hominins Appear?
12
A Note on Geological (& Evolutionary) Chronology
Dates determined by a combination of methods, but especially
Law of super positioning & stratigraphic associations
Potassium-Argon (K-Ar) dating
This Photo by Unknown Author is licensed under CC BY-SA
Feder (2017:54)
Based on principles of radioactive decay, similar to radiocarbon
40K and 40AR contained in fresh volcanic material
AR dissipates when exposed to surface
Over time 40K decays (radioactive and unstable), produces
40AR at constant rate
Half-life of decay process is 1.25 billion years
Huge error range in dates
Also is a predictable relationship between 40AR/39AR
Less error range & more precise

14. 13
(From Feder 2017:27)
We have so many dating techniques for a number of reasons:
Materials we can date
Association of materials we can date with archaeological (and
fossil) record
Applicable range for techniques
Access to (and affordability) of techniques
14
Stratigraphy
Is the first step in all dating because stratigraphy provides
CONTEXT
Good stratigraphic excavation = reliable relative chronological
sequence for the deposition of different layers
This is very important because if something in one deposit is
later dated using an absolute technique, then the whole deposit
can be dated by association
15
Association is important because for archaeologists are not so
much interested in dating the layers or deposits but the
materials that humans have left within them

15. The interconnection of stratigraphic sequences with absol ute
dating methods provides the most reliable basis for dating
archaeological sites and their context
Thanks to the work of Nicolaus Steno (1916) we have several
basic principles of stratigraphy that allow us to use it in dating.
The most important/relevant to this course are:
1) Law of superposition - Layers are successively deposited,
one after another, such that the oldest layers are on the bottom
and the youngest on the top
2) Law of original horizontality - Deposits are laid down
horizontally; Deviations are caused by either uplifting or
downfall
3) Principle of Association - Items found together in same
deposit are of essentially the same age. Caution: humans are
collectors (think heirlooms, treasured tools, which represent
curatorial or archival behaviour. This means that it is possible
to find artifacts that are much older than the context they are
found in.
4) Principle of Reversal - deposits have been removed from site
and redeposited in reverse order; please note that this is rare in
nature but common in cultural contexts
5) Principle of Intrusion - Intrusion must be more recent than
the deposits through which it cuts e.g. underground oven,
garbage pit, well etc.
15
BASIC PRINCIPLES OF STRATIGRAPHY
16
Layers are successively deposited, one after another, such that
the oldest layers are on the bottom and the youngest on the top

16. Law of superposition
Deposits are laid down horizontally
Deviations are caused by either uplifting or downfall
Law of original horizontality
-Steno – Danish Scientist from 17th Century:
Principle of Superposition
-sequence of strata reflects order of deposition
Principle of original horizontality
- strata that are now inclined or even vertical were
originally laid in horizontal position- not rocket science but
relates to concepts of natural transformation processes
STRATIGRAPHY: OTHER PRINCIPLES
17
Principle of Association
Items found together in same deposit are of essentially the same
age
Principle of Reversal
Deposits have been removed from site and redeposited in
reverse order
Principle of Intrusion

17. Intrusion must be more recent than the deposits through which
it cuts
VIDEO: Dr. Biittner explains Stratigraphy and walks through an
exercise showing how these principles are applied
https://youtu.be/gJ5CyyLSmKI
17
14C (RADIOCARBON) DATING
BACKGROUND:
Libby
1947 (1950’s)
Uses organic material
You CANNOT date fossils using 14C dating because during
fossilization the organic materials are replaced with
minerals…no carbon to date!
Half-life 5568±30 years
Generally accepted = 5730±40 years
Three isotopes of carbon
12 & 13 are stable, 14 is radioactive
14C is product of flux of cosmic rays in atmosphere (reaction of
neutrons with Nitrogen 14)
18
Radiocarbon dating (14C) is the single most important method

18. of dating in archaeology.
Based on the ability to measure the rate of radioactive decay.
Used for ages up to 50,000 years old.
Developed by Willard Libby in 1949 which is why 1950 is used
as date from which prehistory is measured. Thus B.P. means
years before 1950.
14C dates using half-life of 5730 years. Means 50% of the
remaining 14C is lost every 5730 years.
18
14C (RADIOCARBON) DATING CONT.
During life of organism 14C is constantly replenished
Exchange ceases with death
Proportion of 14C to stable C will decay logarithmically
(exponentially)
Measurement of 14C relates to the time organism ceased
exchanging carbon with the biosphere
= time of its death
19
19
THE CARBON CYCLE & RADIOCARBON HALF-LIFE
Watch: WATCH: Radiometric decay (half-life) video
https://www.youtube.com/watch?v=-
KYYsknr2Ko&list=PLfpP1vMOJhHWPyPUkHiXlCbnbFQY3IGi
Y&index=8

19. Note that other “radioactive clock” methods like K-Ar and Ar-
Ar work using the same principles – just different materials and
isotopes are used. The effective time ranges of these radioactive
dating methods reflects the varying half-lives of the isotopes
involved.
20
14C (RADIOCARBON) DATING CONT.
Assumes that there has been a constant amount of 14C in the
atmosphere over time
This is false so we have correction curves to calibrate our dates
Assumes that the amount of 14C in the dated material remained
constants since deposition
This can also be false because of contamination
21
21
Note:
mya = million years ago
kya = thousand years ago
Who were the first hominins?

20. Now that we understand some of the background, we can look at
some of the key species.
The final exam study guide has a section on hominins to assist
you with this topic. It outlines again the general information
you need to know (species names, time period, and key traits).
22
How are fossil skeletons studied?
A methodological side bar
Things we ask about the individual
Indicators of:
Age
Stature
Sex
Health
We use this information to create osteobiographies for
individuals
Based on comparison to known range of variation in modern
human populations

21. What are Paleoanthropologists looking for as they classify
fossils taxonomically?
Question for you: Why is classification so difficult?
Should this be considered a hominin?
Based mostly on traits related to bipedalism
Should this be a new genus or species
OR can I fit it in an existing category?
What do variations mean?
What was the lived experience of this individual?
Key Early hominin Sites

22. New finds are constantly changing what is known about early
hominins
Identifying hominins is not as big a problem as deciding which
primate line they are ancestral to
Mosaics of hominin and ape/chimpanzee traits are common
25
Sahelanthropus tchadensis
6 – 7 mya
Toros-Menalla, Chad
Partial cranium, no postcranial
Bipedal?
Closest common ancestor to chimps?
26
What we know right now, the earliest known hominin might be
this guy Sahelanthropus tchadensis
All we have of him is a cranium, which was discovered in 2001
at a site called Toros-Menalla in Northern Chad; unfortunately,
no postcranial bones have been recovered
This fossil hominin has been dated to approximately 7 mya.
The morphology of this hominin is unusual, there is a
combination of characteristics unlike that found in other early
hominins:
Hominid traits include a foramen magnum that is more
anteriorly placed – this is suggestive that the chad may have
been bipedal; his molar enamel is somewhere in between that of
modern humans and of apes, it also has a reduced upper canine
and no diastema.
Apelike features include a small brain and pronounced crests
on the skull where muscles are attaching (so was massively

23. built with huge browridges)
26
5.8 – 6 mya
Western Kenya
19 specimens of jaw, teeth, finger, arm, and leg bones
Orrorin tugenensis
27
-Dating to around 6mya, the fossils of at least five individuals
from this species have been found in Tugen Hills on the western
side of Kenya’s lake Turkana. These finds include portions of
the mandible, teeth, finger, arm, and leg bones. The discoverers
name these hominids Orrorin tugenensis, the genus means,
original man. Among the 19 remains were several partial
femurs, each missing the knee but indicating that these
hominins were bipedal. The molars have thick enamel, like later
hominins.
However, the remains of this hominid does exhibit some ape
like traits. For example, a hand phalanx found at the site is
curved, like a living ape. Also, the upper canine of Orrorin is
large and apelike, throwing a shed of doubt on its hominin
status.
27
The genus Ardipithecus
Includes near complete skeleton “Ardi”
Lived in lightly wooded environment

24. Key traits:
Bipedal traits of pelvis and foramen magnum
Very short thumb (not fully opposable precision grip)
Divergent big toe
Not suited to human gait
Good for climbing
Question: should Sahelanthropus and Orrorin be considered
Ardipithecus?
In 1994 Tim White and a crew of others found Ardipithecus
ramidus in Ethiopia; Publication of White’s finds was delayed
by 17 years to allow for sample preparation and conservation,
and a complete study of the finds.
Ardi has thinner enamel but more ape-like molars. The enamel
on Ar. ramidus teeth remains show it was neither very thick nor
very thin. If the enamel was thick, it would mean Ar.
ramidus ate tough, abrasive foods. If the enamel was thin, this
would suggest Ar. ramidus ate softer foods such as fruit.
Instead, A. ramidus has an enamel thickness between a
chimpanzee’s and later Australopithecus orHomo species,
suggesting a mixed diet. However, the wear pattern and incisor
sizes indicate Ar. ramidus was not a
specialized frugivore (fruit-eater). Ar. ramidus probably also
avoided tough foods, as they did not have the heavy chewing
specializations of laterAustralopithecus species.
This is interesting as well because the first fossil found of Ardi
was a mandibular fragment (some disputed White’s new species
designation) but now have over 100 specimens.
28

25. Ardi’s Foot
29
Limb and foot adapted for bipedalism
Elongated toes
Divergent big toe
29
Ardi’s Place
30
This image shows the likely relationship of the genus
Ardipithecus to the genus Pan (chimpanzees) and the CLCA
(chimpanzee last common ancestor). Note that the genus Gorilla
broke off even earlier from both Pan and Ardipithecus
30
The Australopithecines
Diverse group of early Hominins:
Dates range from 4.2 to 1 mya
Brain not much larger than modern apes
Variation in robusticity
Gracile vs. robust
Bipedal

26. Pelvis and femur
Footprints (3.6 mya, Laetoli, Tanzania)
Partially arboreal
Longer arms than legs, curved fingers
31
-Australopithecus means southern ape
-the genus was named back in the 1920s by Raymond Dart when
he found one
-the australopithecines are represented by hundreds of fossils of
at least seven species
-their fossils have been found mostly in southern and eastern
Africa, in countries such as Ethiopia, Tanzania, and Kenya, and
Africa
-so, they haven’t been found outside of africa
-compared with other mammals, australopithecines do not vary
greatly
-their variation was mostly in size and robusticity – from
relatively small and gracile to large and robust
-some have even gone as far as saying that the two groups,
gracile and robust are so different that they are representative of
two species , australopithecines and paranthropus

27. -as a group, the australopithecines had a small brain, not much
larger than a modern ape
-they were definitely bipedal but some of their appendages show
a potential adaptation to climbing
-they had small canines, large premolars, and large molars
-the later australopithecines (the robusts) had large faces, jaws,
and teeth
31
Australopithecus vs. Paranthropus
A. anamensis
A. afarensis
A. africanus
A. garhi
A. bahrelghezali
A. sediba
A./P. robustus
A./P. boisei
A./P. aethiopicus
Gracile
Robust (Paranthropus)
32
researchers have divided the entire genus up into gracile and
robust forms

28. -the gracile species, also dated to earlier in the fossil record are
the earliest group of australopithecines
-they are usually differentiated form the robust group by their
lighter dentition and smaller faces
-those species included in the gracile group are:
-anamensis
-afarensis
-africanus
-there are others, but these are the most well-known
32
Australopithecus anamensis
4.2 – 3.9 mya
Kenya
-anam: “Lake” in Turkana language
Mosaic of ape and Hominin traits
Most primitive of the genus
33
These are the earliest and most primitive members of the genus
Australopithecus. They lived between 4.2 and 3.9 mya. Anam

29. means lake in the Turkana language; they are named and studied
by American Paleoanthropologist Meave Leakey’s team.
Remains of this species are known from two localities in Kenya.
Mostly known from teeth, but some cranial, mandible, and limb
fragments. With regards to habitat they are thought to have
lived in wooded settings
Anamensis shows a mosaic of traits linking to them to the
earlier Miocene apes and afarensis. They do have some more
hominin-like features: relatively reduced canines, thick enamel
on the molars, indicating selection for grinding hard seeds, the
knee joint suggests bipedalism, and they also have an elbow
joint is like ours, rather than like apes that use arms for support
on the ground.
With regards to their more apelike characteristics: mainly
dental ( parallel tooth rows and a have CP3 honing complex) but
they also have curved phalanges, showing some residual
adaptation to climbing.
33
Australopithecus afarensis
34
3.9 – 2.9 mya
Ethiopia and Tanzania
Most well-known early Hominin:
Lucy (AL-288)
Dikika child

30. A. afarensis is the best known of all the early hominins. Afar is
the name of the local tribe where the fossils were found. It was
discovered in the 1970s in both Ethiopia and Tanzania
Key features of Australopithecus afarensis include ( a) a small
cranial capacity and cranial crests, ( b) a shallow, U- shaped
palate with reduced canine, ( c) features of the postcranial
skeleton that indicate bipedality, and ( d) apelike limb
proportions. They also exhibit sexual dimorphism.
Most researchers feel as thought it was afarensis that
eventually evolved into Homo species.
More than 300 individuals represented by the fossil record –
most famous = Lucy and Dikika child.
34
Australopithecus afarensis
35
Laetoli footprints
Tanzania
Mary Leakey & team
1 to 2 adults and one juvenile
Evidence of bipedalism
Evidence of sexual dimorphism
-Dozens of fossils representing numerous individuals have been
discovered since Lucy, including the footprint trail left behind
in volcanic ash at Laetoli in Tanzania

31. -found by Mary Leakey and her team
-a volcano erupted and covered the ground with ash
-one or two adults and a juvenile walked across the ash just as a
light rain fell
-as the ash dried, it hardened and preserved their footprints
-the prints were clearly made by efficient bipeds, virtually
indistinguishable from footprints left by modern small humans
-what creature made the footprints?
-although some experts argued that A. afarensis was not built to
walk in exactly the way the footprints were made, and held out
for another, yet unknown species
35
Australopithecus africanus
3 – 2 mya
South Africa (Taung, Sterkfontein, Makapansgat)
Bipedal
More human-like dentition
Taung child
36

32. Australopithecus africanus was found in both South and East
Africa. Evidence suggests it lived in open grasslands. There are
no finds of stone tools in the same times and places as A.
Africanus. The body generally like other australopithecines;
small, fully bipedal; although they still have an apelike arm/leg
ratio (arms are longer although the phalanges are not curved).
The cranium is more human-like than earlier australopithecines
or robusts. They exhibit strong sexual dimorphism.
The first australopitheci ne ever found was the type specimen for
the africanus line. A child skull discovered in south Africa, at
the site of Taung by Raymond Dart. This is the type specimen
for the africanus line and because is the face of a 3-4 year old
child (with endocast) it is called the Taung Child. Dart was not
taken seriously when he published in 1925, so he enlisted help
to find more evidence.
Robert Broom was a physician and part-time fossil hunter. At
the Sterkfontein limestone quarry, he found the skull and
endocast of an adult australopithecine. A second site,
Swartkans, yielded more material.
36
Australopithecus sediba
1.95 – 1.78 mya
South Africa
2 partial skeletons
Descended from A. africanus
Derived Homo-like morphology
Ancestral to Homo

33. 37
-evidence for A. sediba was found in South Africa in 2010
-Lee Berger led the team
-the fossils date to 1.97 to 1.78 mya
-Adult and juvenile material was found
-like other australopithecines, the brain size is small, around
420 – 435 cc
-a sediba had long arms – an apelike characteristic
-but, it also has Homo-like traits dental size and shape
-a derived face and pelvis
37
Australopithecus bahrelghazali
3.5 – 3 mya
Chad
First hominin found outside of East African Rift Valley
Partial fragmentary jaw
38

34. -other than a. anamensis and afarensis there are other recently
found australopithecines
-Australopithecus bahrelghazali
-A. bahrelghazali dates to 3.5 – 3.0 mya
-it was found in West Africa
-it is represented by only a portion of a mandible and a few
teeth
-so, from this they created a new species
38
Australopithecus garhi
2.5 mya
Middle Awash, Ethiopia
Few skull and limb fragments
Similar to, but definitely not, any of other australopithecines
Range of variation? Adaptation?
39
-Australopithecus garhi is another of the australopithecines

35. -they date to 2.5 mya
-fossil evidence was found for this species in Ethiopia
-this species had a small brain, around 450 cc
-it had apelike features, it was prognathic and had large canines
-but also had some human characteristics in limb proportions
39
Australopithecus aethiopicus
2.6 – 2.3 mya
Northern Kenya and southern Ethiopia
Ancestral to A. boisei
Hyper-robust
Black skull
(KNM-WT 17000)
2.5 mya
40
Australopithecus aethiopicus lived sometimes between 2.7-2.5
mya ; in East Africa
-this is the earliest of the robusts, because of this, it has some
primitive features, but in many ways the most exaggerated of all
robusts
-it is hyper robust: most massive temporal muscles (flaring
zygomatics) and sagittal crest of any australopithecine but more

36. prognathic (protruding face) – not dished which is less suited to
extreme pressures on the back teeth so maybe a somewhat
different dietary specialization
-small brain, 410 cc
40
Australopithecus boisei
2.3 – 1.1 mya
East Africa (Olduvai Gorge, Tanzania)
Louis and Mary Leakey
Zinj or Nutcracker Man
Tool maker ?
Pronounced sexual dimorphism
41
The second of the robust australopithecines, Australopithecus
boisei lived sometime between 2.3 to 1.2 mya in East Africa. It
is basically an even more exaggeratedly robust version of A.
Robustus: even bigger molars, larger temporal muscles, more
vertical face, etc. All clearly worked to apply and withstand
huge pressures on those giant molars
Males weighed 68kg (150lbs) and stood 1.3m (4’3”); females
weighed 45kg (100lbs) and were 1.05m (3’5”)
This is the greatest sexual dimorphism of any hominin.

37. 41
Australopithecus robustus
2.8 – 1 mya
South Africa
Robert Broom
Omnivore
Tools ?
42
-Australopithecus robustus, obviously one of the robust group,
was around from 2 to 1.5 mya. Their remains have been found
in South Africa.
From the neck down, robustus was not very different from the
other australopithecines. The head, though, is drastically altered
by its massive chewing apparatus (huge molars and premolars,
huge mandible, huge temporal muscles - the main muscles that
squeeze the jaws together). These all create a wide dished out
face. Attachment markings that show the temporal muscles di d
not just attach at the side of the cranium, but covered the whole
cranium, meeting in the middle; this forms a sagittal crest, or
vertical ridge of bone along the top of the head where the
temporal muscles attached. This is all generally assumed to
indicate that A. robustus ate hard or tough plant material and
tooth wear suggests hard seeds or nuts.
Robustus may have made and used tools! Animal bone shafts
with heavy wear on one end have been found at a site where the
only hominin fossils are A. robustus. The wear on the fossil
bones best matches the wear caused by digging anthills and
remember, A. robustus is not considered our ancestor. This
suggests that tool making and use at a level slightly beyond

38. what chimps do developed in at least two different li neages of
hominins: robust australopithecines and whatever led to us.
42
43
Image showing relationships between Ardipithecus,
Australopithecus, and Homo
43
Discussion
Why were there so many Australopithecines?
Which one(s) led to the genus Homo?
“Hmm… That’s a real head scratcher”
The Pleistocene:
What environmental pressures shaped human evolution?
Cooling towards “Ice Ages” begins 2.6 mya ends perhaps with
Holocene 10 kya
Climatic fluctuations:
Glacials – ice caps advance
Interglacials – climatic amelioration, ice retreats
Stadials and interstadials, shorter periods of advance and retreat

39. within glacials or interglacials
Overall effects:
Extreme cold in north
Fluctuations in precipitation and air flow – changes
biogeoclimatic zones, esp vegetation
Sea levels drop
Timing of climatic shifts perhaps correlated with speciation
Or at least a new species had to learn to adapt
Feder (2017:118)
What do we know about early Homo?
46
47
Overview of Hominins and Stone Tool Industries

40. Early Hominin Tools
First tools recovered in East Africa
Earliest dated = Lomekwian (Lomekwi, Kenya) = 3.3 mya
Earliest Oldowan = Gona, Ethiopia 2.5 mya
Most date between 2.5 – 1.7 mya
Australopithecines?
early Homo?
Lower Paleolithic
Choppers and scrapers
48
Tools from Lomekwi
Feder (2017:84)
Before we talk about the early members of the Genus Homo, we
need to discuss something that is very important. At very
roughly the same time as early Homo appeared, the first
recognizable stone tools began to be made – with the emergence
of Homo we have the emergence of Culture.
These early tools, dating as early at 2.5 mya, are called the
"Oldowan" (or Lower Palaeolithic)
They are named after Olduvai Gorge in Tanzania where they
were found (recall photos from last lecture).
They are extremely simple: generally rounded cobbles or cores
with a few flakes broken off to produce a sharp edge. The flakes
were probably used for cutting, too.
Now these tools may have been made by early Homo, although

41. it is virtually impossible to prove that, since australopithecines
were around, too
In fact, the first A. boisei fossil was found in a scatter of
Oldowan tools (at the FLK Zinj site in Olduvai Gorge)
48
49
Oldowan Tools
But what does finding tools and bones in the same place really
mean?
Why would the maker of the tools drop dead right there amid
his or her handywork?
Did this A. boisei use the tools on the carcasses of other
animals found there - or were the tools used on A. boisei?
A. gahri (one mentioned in your textbook) is the only hominid
found in the region of Bouri, where animal bones were found
with cutmarks on them
Did this gracile australopithecines make the tools, or was it
some other hominid that has not been found yet?
49

42. Early Homo: Variability
50
2.4 – 1.5 mya
East and South Africa
Two (2) contemporaneous species?
Homo habilis
Smaller body and brain size
Gracile face and teeth
Homo rudolfensis
Larger body and brain size
Robust face and teeth, more similar to Australopithecines
Homo habilis is the first member of our genus. At least for now.
It dates to between 2.5 and 1.7 mya. It was named by Louis
Leakey, Philip Tobias, and John Napier. Homo habilis means
“handy man” – anyone want to guess the significance of this
name. Homo habilis is known from Tanzania, Kenya, Ethiopia,
Malawi, and South Africa, which is the same geographic
distribution as the Australopithecines
Brain size:
> brain than Australopithecines
50% less capacity than modern humans
Dentition
More similar to later Homo
Cranial anatomy
More rounded, more similar to Homo
Postcranial anatomy
Fully bipedal (?)

43. -examples: OH 7, KNM-ER 1813
Homo rudolfensis
-H. rudolfensis is a more heavily built species (with more
human like limb proportions)
-they have larger teeth
-a more massive, wide, sloping face
-and a considerably larger brain, maybe around 775 cc
-example: KNM-ER 1470
Sexual dimorphism or separate species ? ? ?
50
Homo naledi
Rising Star Cave, South Africa
Over 1500 bones
Minimum number of individuals (MNI) = 15
Date between 335 – 236 kya (!!!)
Key Traits
A mosaic of traits:
Smallish brain
Curved fingers
“modern” foot
How did they get in there?
Homo naledi: Rising Star Cave

44. Who was Homo erectus?
53
1.8 mya – 500 kya
As late as 100 kya in Asia
Great diversity/variability anatomically
Just H. erectus or H. erectus AND H. ergaster
Africa: Olduvai Gorge, Tanzania; Lake Turkana, Kenya;
Swartkrans, South Africa; Omo, Ethiopia
First movement out of Africa (Out of Africa I)
Trinil and Sangiran, Java
Swanscombe, England
Zhoukoudian, China
Sima de los Huesos and Gran Dolina, Atapuerca, Spain = Homo
antecessor
Dmanisi, Georgia (W. Europe) = Homo georgicus
When specimens of homo erectus were first found in 1891 by
Eugene Dubois, he called them Pithecanthropus erectus, or erect
ape man - we now refer to this hominid as Homo erectus. For
the first 5 million years of evolution, hominids had been
restricted to Africa, but this species quickly found its way out
of Africa, traveling as far as Southeast Asia, discovering new
tools, foods, and technology. Like Homo habilis, there some
morphological variation among the different geographical

45. groups of these highly successful hominids because of this,
palaeoanthropologists are still debating how to classify them
(we’ll get to this later). Homo erectus populations lived far
across the world from one another; they are however, united by
common traits.
The skull of homo erectus has some characteristic traits. Some
of them are more primitive, meaning they’ve retained them from
early homo, and others are more derived, meaning they’ve
developed them since the split from homo
53
Homo erectus
54
Despite diversity, some typical traits can be identified in the
cranium & postcranium
Cranium
Pronounced ridges, sloping forehead, receding chin
Smaller more vertical face
Smaller teeth
Postcranium more similar to modern humans than to early Homo
Larger body size
Modern proportions
The primitive characteristics retained from early Homo are:
solid, bar-like browridges (supraorbital torus), low, sloping

46. forehead, and receding chin, so, no real chin like ours (later,
you will see that having a chin is a human characteristic),
pronounced muscle attachment areas on the back of the
skull.These traits could indicate a reduced emphasis on
grinding, and a greater emphasis on holding, cutting, and
tearing with the front teeth. The molars are the teeth that are
most reduced. The occipital torus probably indicates stronger
muscles at the back of the neck, which would be needed to use
the teeth for pulling and the massive browridges may strengthen
the face to resist the forces involved in pulling with the front
teeth.
Their derived characteristics, or those found in modern humans
include: more vertical face (less prognathic), smaller face (not
as tall), and smaller teeth, especially the molars which is a
major reduction in the back-teeth grinding adaptation of earlier
hominins. H erectus is apparently the first hominid to have a
projecting, human-like nose; this might reduce evaporation,
conserving water in dry climates or it might provide a radiator
to help cool blood to the brain
In general, the postcranium (everything but the head) of Homo
erectus is very similar to modern humans; well at least,
definitely more similar to us than it is to early homos like
habilis. Erectus has less sexual dimorphism than early Homo;
about the same as modern humans; male H. erectus were only
20% to 30% bigger than females (Today in the US, males
average about 25% heavier than females). It is thought to have
limb proportions like us, meaning that their legs are longer than
their arms, making them well-suited to bipedalism and giving
them a nice striding gait.
54

47. Nariokotome (Turkana) Boy
KNM – WT 15000
Nearly complete H. ergaster/erectus specimen
Dates to 1.6 my
Sex = Boy = shape of pelvis
Age = 11 – 13 = dental eruption
(no 3rd molar = no wisdom teeth)
Skeleton = 5’3”
Projected would have stood between 5’6” and 6 feet tall had he
reached maturity
-the amazing find of KNM-WT 15000 (the "Nariokotome boy")
-a nearly complete skeleton of an Homo ergaster boy of about
12 years old who was already 5 feet 6 inches tall!
-he dates to about 1.6 mya and is very complete, gives us a good
luck at the species’ postcranial skeleton and even the maturation
of the skeleton
55
Homo erectus: Culture

48. 56
Acheulian Tool Tradition
Earliest finds in East Africa (1.5 mya)
Latest finds vary: Europe (~100 kya)
Big Game Eating
Control of Fire
Schöningen, Germany
Language?
Other ritual evidence
1. Homo erectus continued to use the Oldowan chopper tool
technology but also made better stone tools than did early
Homo. Starting around 1.4 mya in Africa, H. erectus also made
more elaborate tools we call "Acheulean“. Acheulean tools
typically have flakes knocked off from both sides to form a
bifacial edge but instead of forming just a point or short edge,
the edge goes all the way around the tool making a bifaces and
instead of being shaped like the original cobble with a few
flakes banged off, the tool has a distinct, intentional form. The
most typical form is the handaxe, which is fairly flat and
teardrop shaped. These regular, consistent forms indicate that
they were intentionally made that way and that H. erectus had
the mental ability to visualize the shape of the tool, and the
skill to realize the plan = mental models. There is some debate
about what these Acheulean handaxes were used for. Some of
the more reasonable possibilities that have been suggested: used
like axes to chop up carcasses of large animals? used for
digging for roots, burrowing prey, and/or water fowl hunting?!

49. Importance of colour, placement of fossil
2. Some of the Acheulian sites have produced evidence of big-
game eating. For example in Spain they found a substantial
number of elephant remains and unmistakable evidence of
human presence in the form of tools but, so these finds of bones
in association with tools tell us that the humans definitely were
big-game hunters?? Some think the big game may have been
scavenged. It is still unclear whether erectus deliberately
butchered different kinds of game; although unclear how they
killed them, we can be sure that they consumed it and most
definitely hunted smaller game. Other sites have possible
evidence of communal hunting, where animals may have been
driven into a specific area.
2. & 3. At Schoningen, Germany archaeologists have recovered
400 ky old spruce spears that are fire hardened along with other
evidence for systematic hunting. Evidence from Boxgrove,
England and Torralba and Ambrona sites in Spain also suggest
cooperative hunting.
3. Further because erectus was the first hominid to be found
throughout the old world and in areas with freezing
temperatures, most anthros believe they had learned to control
fire. There is archaeological evidence of fire in some early
sites, but fires can be natural events thus, whether fire was
under deliberate control by erectus is difficult to establish. It is
hard to imagine that erectus could move into these areas without
the use of fire, and if not, clothing would have been necessary.
4. We can use skeletal evidence to determine whether homo
erectus had developed language. With erectus, there was a
remodelling of the brain, in particular differential growth of the
frontal and posterior portions of the brain; in other words, homo
erectus brain is more similar in form to our brain. Analysis of
the endocast, or the fossilized brain, of some erectus skeletons
have showing that their right and left halves of the brain control
different aspects of their behaviour, just like us; this has been
interpreted by some to suggest that by 1.7 mya erectus would

50. have possessed linguistic skills and the ability to manipulate
symbols. Some have suggested that it had the language of a 6
year old; however, other research has shown the exact opposite
- that erectus’ brain development is more like that of apes than
modern humans. Thus the jury is still out on this one.
5. Bodo site in Ethiopia – cut marks around eye socket on
cranium; Possible ritual defleshing of cranium; At Sima de los
Huesos (Atapuerca, Spain) argued see deliberate, ritual burial;
finally, possible female figure in volcanic scoria at acheulian
site of Berekhat Ram (Israel) dating to 230,000 ya
The archaeological finds of tools and other cultural artifacts
dating from 1. 8 mya to about 200,000 ya are assumed to have
been produced by Homo erectus
Unfortunately, fossils are not usually associated with theses
materials; therefore, it is possible that some of the tools were
produced by hominids other than erectus such as habilis or even
later homos
With regards to erectus’ culture, we can say a few things about
their tool tradition, diet, use of fire, and language
Other Ritual Evidence
Bodo site in Ethiopia
Cut marks around eye socket on cranium
Possible ritual defleshing of cranium
Sima de los Huesos, Atapuerca, Spain
Deliberate, ritual burial
Berekhat Ram, Israel
Possible female figure in volcanic scoria
230 kya (late Acheulian)
56

51. Isimila, Tanzania
57
Comparisons of estimated cranial capacity (cc) & overall
cranial morphology
58

52. The “Hobbit”: Homo floresiensis
Discovered in Liang Bua Cave on Indonesian island of Flores
(2003)
~1m (3ft) tall
Fully bipedal
Cranial capacity ~380 cc
Fossils date: 100-60kya
Archaeological dates: 190-50 kya
Similar to that found with H. erectus species
59
We also have to deal with the coolest dude ever, homo
floresiensis
Whether or not there are distinct African and Asian species of
erectus, some have suggested that Homo floresiensis is a
distinct species that is closely related to erectus
Homo floresiensis has only been found on the Indonesian island
of Flores, and only a handful of individuals have been located
to date
All are tiny, they stood perhaps just under one metre tall, or
around 3 ft
They had very small brains, on the order of 380 cc

53. How might a miniature version of H. erectus have evolved? The
answer is that both dwarfism and gigantism are common
phenomena in isolated populations. Dwarfism appears to be an
adaptation that occurs when there are few predators; a species
in isolation can become smaller, and have a large population, if
there are no predators threatening them, and this may be what
happened on Flores. The island also hosted a number of dwarf
species in addition to Homo Floresiensis including a dwarf
elephant that appears to have been one of the favourite foods of
floresiensis
However, others contend that a disease process could explain
the smallness of floresiensis. Pathological conditions such as
microcephaly, a developmental defect, would explain the small
cranium
Even more interesting is that floresiensis may have survived
until as recently as 12,000 years ago, well into the time period
when modern humans were living in Indonesia
Could modern humans and the time floresiensis have met some
time in the distant past?
We do not know as there is no evidence
59
How did Homo sapiens evolve?

54. Like H. erectus, premodern hominids were spread throughout
the Old World
Europe was permanently and in some areas, densely, populated
Their range was not much greater than that of H. erectus whom
they replaced
60
Transition to Modern Humans: Context
Archaic H. sapiens
First appear in the Middle Pleistocene (900–125 kya)
Some persist into the Late Pleistocene (125–10 kya)
Pleistocene? (aka Ice Age)
Glacial: Ice cover in Northern Eurasia; Africa is dry
Interglacial: No ice cover in Eurasia; Africa Wet
Hominins affected by changes in climate, fauna, and flora
61
61
Archaic Homo sapiens
Also called ‘Homo heidelbergensis’
Found in Africa, Asia, and Europe
Did not vastly extend the geographic range of Homo erectus
Some difficult to distinguish from Homo erectus
Key traits include:
Larger brain

55. More globular brain case
Arching, heavy supraorbital torus
Occipital torus
Low forehead
Smaller teeth and jaw, larger brain, smaller more gracile skull
than H. erectus
Large, prognathic face and teeth, brow ridge, low, rounded skull
compared to H. sapiens
62
Variation:
The fossils from Europe and Africa are closer in appearance to
one another than to those from Asia – they are usually given the
name H. heidelbergensis
The situation in Asia is less clear – some fossils appear more
modern than those from Europe and Africa
Glaciation isolated populations and so regional variations could
have easily arisen
In Europe, H. heidelbergensis may have evolved into
Neanderthals, while the Chinese premodern populations may
have gone extinct
62
Archaic Homo sapiens: Culture
Acheulian tool tradition continues

56. Some later groups used the Levallois technique (Middle Stone
Age/Middle Paleolithic culture)
Several changes to more ‘modern’ culture seen:
Apparent use of fire
Simple shelters
Wider range of foods
Large animals hunted
63
63
Who were the Neandertals?
64
Neandertals & Pop Culture
65
Rosny, Quest for Fire, 1911 (1981)
HG Wells, The Grisly Folk, 1927

57. William Golding, The Inheritors, 1955
John Darnton, Neanderthal, 1996
Our fascination with this species so similar and yet so unalike
us is evident in popular culture: they are the topic of choice in
film and novels, tv specials, comics and more!
Neanderthal = thrilling new york times best seller!!
Clan of the Cave Bear series by Jean Auel (6 books!!!!)
Unfortunately many of these popular depictions are more fiction
than fact.
So who were the Neandertals?
Who were the Neandertals?
Late European/West Asian premoderns
Named after the Neander Valley
Stem from earlier premoderns
First definite Neandertal remains date to 175 kya
Biache-Saint-Vaast, France
Latest Neandertal remains date to 27 kya
Mezmaiskaya Cave, Russia
66

58. First found 1856
350 – 30 kya : during the Pleistocene (aka the ice age = period
of great climatic change= floral and faunal change)
the Neandertals are some of the best-known and most-studied
fossil hominids in the world
-owing to the relative completeness of the fossil record,
palaeoanthropologists have been able to document and debate
the meaning of their physical characteristics
-since their discovery more than a century ago, Neandertals
have haunted the minds and foiled the best laid theories of
paleoanths
-there is disagreement on how to classify them, which is why
they have several names
-considerable controversy continues among researchers as to
whether or not the Neandertals were ancestral, in whole or in
part, to modern humans
those who do not believe the Neandertals were ancestral,
essentially resulting in a dead end, refer to the Neandertals with
a separate species designation
-however, many others believe that the Neandertals were at
least partially ancestral to modern humans and separate them
only by subspecies designation – homo sapiens neanderthalensis
66

59. 67
Neanderthal Anatomy
-Neandertal morphology is quite distinctive, especially in the
skull
-one striking feature of Neandertals is brain size which in these
hominids was larger than that of homo sapiens today; average
cranial capacity for modern humans between 1300 and 1400 cc
where the average cranial capacity for Neandertals was
somewhere ~1600 cc
-the classic Neandertal cranium is large, long, low, and bulging
at the sides; viewed from the side, the occipital bone is
somewhat bun-shaped; researchers have hypothesized that this
bun may have developed to keep the head stabilized during
running but really, they have no idea why it is there
-the forehead rises more vertically than that of homo erectus;
similar to archaics, their brow ridges arch over the orbits
instead of forming a straight bar as in erectus
-compared with anatomically modern humans, the Neandertal
face stands out; it projects almost as if it were pulled forward
-they are also characterized by other diagnostic traits such as
the juxtamastoid eminence and the retromolar space
-finally, they have a very large nasal aperture, meaning that
their nose was pretty big
Similar to modern humans in many ways
Key differences include:
Barrel-chested

60. Shorter limbs
Generally more robust skeleton (hyper-robust)
More powerfully muscled
67
Neandertals: Adapted for Cold?
Neandertal cranial & postcranial morphology has also been
interpreted as an adaptation to cold climate
Body:
Short and wide, shortened limbs
Build may reflect need to conserve heat
Facial morphology:
Large brain
Large nasal aperture
Midfacial projection
Infraorbital foramina
68
-Most distinctive about the cold adaptation complex in
Neandertals are the shape of the body trunk and the length of
the arms and legs; compared with modern humans, European
Neandertals were short, wide, and deep and their limbs were
shorter
-this combination, stocky trunk and short limbs is predicted by
Bergmann’s and Allen’s rules that is, animals that live in cold
climates are larger than animals that live in hot climates
(Bergmann’s rule). the larger body trunk reduces the amount of
surface area relative to the body size this helps promote heat
retention

61. -finally, animals that live in cold climates have shorter limbs
than animals that live in hot climates (Allen’s rule). this too
promotes heat retention in cold settings. so, the shorter distal
limb segments (forearm and lower leg) and barrel chest in
neandertals decreased the surface area to volume ratio of the
body in order to retain heat
because Neandertals lived during glacial times, researchers feel
that the larger brain size may be associated with the metabolic
efficiency of a larger brain in cold weather
-because of their enormous nasal aperture, researchers have
argued that they had a big nose, in both width and projection:
one of the noses’ important functions is to transform the air
breathed in to warm humid air; arge noses provided more
internal surface area, thus providing an improved means of
warming and moistening the cold dry air
-moreover, the projecting nose, because of the midfacial
projection, placed more distance between the cold external air
and the brain which is temperature sensitive
-other features of the Neandertal skull that are consistent with
cold adaptations include the infraorbital foramen. these are
small holes in the maxilla found under the orbits. well, they are
exceptionally large in European Neandertals compared to
modern people. the foramina’s increased size is due to the blood
vessels that tracked through them having been quite large. the
larger blood vessels allowed for greater blood flow to the face,
preventing exposed facial surfaces from freezing
These ideas were are not excepted by everyone however.
68

62. Neandertal Anatomy
Strenuous
Show many (healed) injuries
Neandertals show injury pattern similar to modern rodeo riders
Proposed by Thomas Berger and Erik Trinkaus
May reflect more need for strength compared to recent humans
May reflect close approach to large prey (i.e., hunting strategy)
69
Valerius Gerst (2000) developed the bull riding hypothesis
based on an impression drawn from visit to the Calgary
Steampede while thinking about differences in Cro-Magnon
(UP) and Neandertals (Mousterian) archaeological records:
Neandertals focused on larger prey (prey with long hair perfect
for clinging on like woolly mammoths, woolly rhinos & horses;
while one hunter would distract it another would cling onto it
(the distractor could then get in closer to bucking animal and
kill it)
-two American anthropologists, Thomas Berger and Erik
Trinkaus studied Neandertal trauma patterns in order to
understand a societies behaviour (associated risks); the
researchers noticed that nearly every complete Neandertal
skeleton displays some traumatic injury
-they compared the patterns of injuries with that of modern day
occupations; interestingly, the Neandertal pattern resembles that
of rodeo athletes (the people who ride angry broncos and bulls
as a form of sport) -rodeo riders have lots of head and neck
injuries resulting from the obvious

63. Does this mean Neandertals climbed trees and jumped on to
their prey?
69
Neandertal Culture: Technology
70
Anthropologists almost always associate Neandertals with
Mousterian technology (180 – 30 kya) although, archaic homo
sapiens were also using it
-the Neandertals improved on previous prepared core techniques
– that is the Levallois by inventing a new variation
-they trimmed a flint nodule around the edges to form a disk-
shaped core
-each time they struck at the edge, they produced a flake and
they kept at it until the core became too small and was
discarded -in this way they produced more flakes per core than
their predecessors
-they then reworked the flakes into various forms including
scrappers, points, and knives
-they elaborated and diversified traditional methods, and there’ s
some indication that they developed specialized tools for
skinning and preparing meat, hunting, woodworking, and

64. hafting
Varied, regionally diverse tool kits = Binford & Bordes Debate
(text)
Compound (hafted) tools
70
Neandertal Culture: Subsistence
2.- subsistence - based on animal remains at several Neandertal
sites, they were successful hunters
-it’s clear that they hunted large mammals but we’re just not
sure how good they were at it
-they had no long distance weaponry, so they were limited to
thrusting spears hence the trauma
Archaeological evidence shows Neanderthals were capable of
hunting large game
At La Cotte de St. Brelade site, at least 20 mammoths and 5
woolly rhinos were deliberately stampeded off a cliff then
butchered by Neanderthals
At Umm el Tlel, a fragment of a Levallois point was found
embedded in a vertebra of a wild ass
Bone chemistry studies show that Neanderthals were essentially
meat eaters—one individual’s diet appears to have been 97%

65. meat
Neanderthals had base camps where resources were brought
back for consumption
Many Neanderthal base camp sites are located in caves
Many of these sites show evidence of intensive occupation and
deep accumulation of archaeological layers
Much of the sediments at these sites are products of human
activities—charcoal, bone fragments, stone tool fragments
Neanderthal cave sites are spatially organized
Main living surface was relatively clean, while the sides of the
cave were refuse areas
In the central living areas were hearths
—not a defined fireplace, just many fires over time
71
Neandertal Culture: Speech
72
Philip Lieberman and Edmund Cremlin reconstructed a
Neandertal vocal tract – not capable of speech
Hyoid found at Kebara, Isreal – capable of speech
Fundamental to human behaviour is the ability to speak
-because early anthros believed that Neandertals lacked the
ability to speak, they believed that Neandertals were not related
to modern people in a evolutionary sense

66. -these ideas persist today
-Linguist Philip Lieberman and Anatomist Edmund Crelin have
reconstructed the Neandertal vocal tract
-because their reconstruction resembles a modern newborn, they
conclude that, like human babies Neandertals could not express
the full range of sounds necessary for speech
-however, another compelling line of evidence suggests that
Neandertals were able to speak
-I’ve already mentioned this but, a Neandertal skull found at the
Kebara site in Isreal included a hyoid
-again, various muscles and ligaments attach it to the skull,
mandible, tongue, larynx, and pharynx
-collectively, these elements produce speech
-the morphology of the kebara hyoid is identical to that of a
living humans
-therefore, the Kebara people talked
72
Neandertal Culture: Burials
Definitely intentional
Le Moustier
La Ferrassie
Shandiar (IV)
Krapina
73

67. -in many Neandertal sites, their remains have been found
scattered around, commingled and concurrent with living areas
-For example, the Krapina Neandertal fossils from Croatia are
fragmentary and were scattered throughout the site that is, the
deceases were treated no differently than food remains or
anything else being discarded
-in contrast, a significant number of skeletons have been found
in pits that is, excavations in Europe and Western Asia has
shown that pits had been dug, corpses had been placed in the
pits, and the pits had been filled in
-For example, the Neandertal skeletons from various sites in
France such as La Chapelle aux Saints, Le Moustier, La
Ferrassie, and nine!! Shanidar individuals from Iraq and some
from Amud and Tabun in Isreal were found in pits
-was burial of the dead a religious or ceremonial activity having
significant meaning for the living? or was the burial simply a
means of removing bodies from living spaces?
-most of the intentionally buried skeletons were in flexed
positions; the hands and arms were carefully positioned and the
bodies were typically on their sides or backs
-this vigilant treatment indicates that care was taken to place
the bodies in the prepared pits so, this pretty much means that
these burials were not just disposals
-even more so, the evidence at Shanidar consists of pollen

68. around and on top of a Neandertal man’s body; pollen analysis
suggest that the flowers included ancestral forms of modern
grape hyacinths, yarrow, cornflower, bachelors buttons (all
known for medicinal properties as diuretics, stimulants,
astringents and antiinflamitorys). was this intentional? Or was
the pollen placed there by humans or was it natural
(bioturbation)?
Neanderthals sometimes buried their dead in small pits,
sometimes with grave goods
At Kebara Cave, the outline of the burial pit can be traced
stratigraphically
At Amud Cave, an infant was found in a natural niche in the
side of the cave with the upper jaw of a red deer
Neanderthal remains from sites in France and Croatia show
evidence of cannibalism
Some of these Neanderthal bones show evidence of butchery in
the form of cut marks from defleshing and percussion marks
from smashing the bones to obtain marrow
73
Neandertal Culture: Care
Healed Injuries:
Spinal, rib, limb fractures
Amputation
Blindness
Degenerative joint diseases (i.e., arthritis)
Teeth loss (bad teeth!)
Little or no signs of infection
Care for elders
Shanidar 1
Elderly man with withered right arm (paralysis?), walked with a
limp, deformity to left side of skull

69. 74
Enamel hypoplasia = indicator of dietary stress
Shanidar I is called Nandy (40 – 50 years old which is like 80
years old today)
Withered right arm from disease or trauma, and survived a
crushing blow to the skull which resulted in blindness
74
Us or Them?
Differing interpretations of placement in human lineage and
intelligence over time
Dumb brutes – flower children – social peers
Neandertal prejudice/racism?
75
They key point of all this discussion of Neandertal
characteristics – relating to climate adaptation, efficiency in
hunting strategies, treatment of the deceased, and use of speech
is that Neandertals likely were not weird human-like primates,
less adaptable and less intelligent than modern humans
-the record shows that their behaviours were humanlike, similar
enough to modern humans

70. -therefore, Neandertals were definitely not less than human as
pervious people have thought
Neandertals as Beasts
Early interpretation (pre-WWII) of Neandertals
1909 by Kupka – rendered from original 1856 fossil specimen
- See beast-like characteristics: very harry, ape like face with
aggressive/unthinking expression, crude club, bones scattered as
debris at feet
76
NEANDERTALS AS US
In the 1990’s and into the 2000’s more and more support has led

71. to the opinion that Neandertals were us (or at least close enough
that their contemporary modern humans would have seen them
as similar enough to comingle with them)
This change in the tide of opinion is directly linked to
Rejection of racism and racist interpretations
Rise of genetic analysis and increasing amount of genetic
evidence
Neandertals & Genetics
Genomes of people living outside of Africa today are composed
of some 1 to 4 % Neanderthal DNA
Influenced skin and hair colour (POU2F3, BNC2)
mtDNA
At least 99.5% identical
Neanderthal-human common ancestor = 706 kya
Neanderthal-human split = 370 kya
nDNA
Genome sequencing (Green et al. 2010)
Evidence of admixture or common ancestor? (Yotova et al.
2011)
FOXP2 (capacity for speech)
MC1R (“Ginger” man?)
Problems = decay, fragmentation, and contamination
Recall that we are 98.5% chimpanzee
Work by Sankararaman and Reich: used sequenced Neanderthal

72. genome and screened 1,004 modern genomes for sequences with
distinctive Neanderthal features. For example, if a fragment of
DNA is shared by Neanderthals and non-Africans but not
Africans or other primates it is likely to be a Neanderthal
heirloom. Also Neanderthal sequences are typically inherited in
large batches since they were imported into the modern human
genome relatively recently and have not had time to break apart.
Some clear influences: the Neanderthal version of the skin gene
POU2F3 is found in around 66% of East Asians while the
Neanderthal version of BNC2 which also effects skin colour is
found in 70% of Europeans – argued helped moderns adapt to
new environments.
Neandertal mitochondrial DNA sequencing provided no
evidence of Neandertal matrilinealcontribution to contemporary
humans. In contrast, a draft
sequence of the Neandertal nuclear DNA revealed the presence
of a number of bona fide Neandertal segments in non-African
genomes (Green et al. 2006, 2010). Yet, although rather
unlikely given all the precautions taken to eliminate such
a possibility (Green et al. 2010), the admixture results could be
due to contamination of ancient DNA with modern human
genomic fragments (Wall and Kim 2007). Yotoval et al. 2011
provide further evidence of Neandertal admixture. Figure on
the right illustrates the plausible historical pathways leading to
the 3 observed categories of dys44 haplotypes represented in a
sample of 6,092 x-chromosomes representing populations from
all habitable continents. Basically, one of the haplotypes,
which carries two types of derived alleles that are shared with
Neandertal DNA, is absent from Africa therefore indicates
1. very early Neandertal admixture prior to successful range
expansion of the population ancestral to virtually all
contemporary non-Africans OR
2. That ancestral population of present-day non-Africans was
more closely related to Neandertals than the ancestral
population of present day Africans

73. FOX2P= provides for finer control over the mouth and throat
which is essential for articulating sound differences;
pronounciation but also word order and comprehension
Not a gene for language but a precondition for human speech =
found that neandertals share the human variation of this gene!
Melanocortin 1 receptor allele (MC1R)
Regulates pigmentation (balance between red-and-yellow &
brown-and-black) in humans and other vertebrates
Living people with mutations that reduce activity of MC1R tend
to have red hair and pale skin
Lalueza-Fox et al. (2007) found a mutation that affects MC1R
in this way in two Neandertal specimens
From: http://www.geneticliteracyproject.org/2015/05/12/more-
mystery-about-neanderthal-and-humans-how-reliable-is-ancient-
dna-analysis/
“the analysis of ancient, partially decayed, and fragmentary
DNA inevitably includes making certain assumptions and
decisions about missing and degraded pieces, in order to fill in
the gaps as best as possible. Some decisions about assembling
Neanderthal sequences are based on sophisticated algorithms
involving the use of modern human and chimpanzee genomes as
guides. Further challenges are presented by the need to filter out
contamination by modern DNA, such as from the researchers
who handled the bones or from microorganisms that lived
within or among the buried bones.
These challenges and complexities raise the possibility of the
inadvertent introduction of errors in the sequencing process —
even when all precautions are taken by highly professional
researchers. These issues also open up any obtained data to
different interpretations and to possible limitations in the
application of the data.”

74. What happened to the Neandertals?
Interbreeding
Hybrid individuals
Fossil evidence
Genetic evidence (mtDNA)
Means did not live as separate, independent group since ~40 kya
BUT might not mean interbreeding
Genocide
No evidence
Extinction
Retreat & refuge
“Poor mother” hypothesis (Paul Mason)
79
According to a model developed by Cambridge evolutionary
ecologists Andrea Manica and Anders Eriksson, “There was an
ancestor of both Neanderthal and modern humans—some
archaeologists would call thatHomo heidelbergensis—that
would have covered Africa and Europe about half a million
years ago.”
In other words, just because some people today apparently share
certain segments of DNA with extinct Neanderthals, that does
not necessarily mean that interbreeding occurred.
Few cases of “stab” wounds: Shanidar II: lung punctured
(probably) by stab wound to chest between 8th and 9th ribs.
Intentional attack or hunting accident? Survived for some weeks
before being killed by rockfall in Shanidar cave. Otherwise is a
sensational story but story nonetheless. Absolutely no direct
evidence.
Idea that neandertals could not compete with modern humans

75. has the best archaeological support: see retreat from Near East
into eastern then Western Europe = can see retreat when look at
dates of sites. Appears to be refugee population in Iberia 30
kya.
www.cosmosmagazine.com/features/better-mothering-defeat-
Neanderthals/
Argument by Paul Mason that human mothers, in comparison to
their Neanderthal sisters, lavished more attention to their
offspring. Essentially evidence to date suggests that
Neanderthals were patrilocal whereas modern humans appear to
more likely have been matrilocal. Female Neanderthals were
physically robust and hunted alongside their men – their fossil
bones show that they sustained injury from close proximity big
game hunting. Humans on the other hand appear to have opted
for a greater division of labour. While female Neanderthals
were off hunting with the men, female humans spent their time
on domestic duties, which included cave painting. This find
highlights the creative role that female humans played in
foraging societies, and also has clear type of mothering they
were able to deliver. Analysis of a tooth from a Neanderthal
infant suggests that Neanderthal children moved off exclusive
breastfeeding at around 7.5 months of age. In comparison,
infants in human foraging societies are not weaned until 3 or 4
years of age. Clearly female participation in close-range hunting
would impact negatively upon child-rearing capacities while
lullabies and cave painting would be more conducive to the
mother-infant bond. Paul Mason argues that the human babies
who had more time with their mothers, who were in turn
supported by their own families in the task of raising highly
dependent, slow maturing, offspring found themselves in an
ideal learning environment. The mother-infant bond is important
in terms of language, nonverbal communication, and
interpersonal emotional skills, therefore extended childhood
increased the duration of learning and innovation.

76. Anatomically Modern Humans
80
Anatomically Modern Homo sapiens
First appear ~200,000 years ago during the Middle Pleistocene
Earliest dated = 180kya at Omo, Ethiopia
By 50,000 years ago they had spread across the globe, even as
far as Australia!
Contemporaneous with Neandertals
81
81
Anatomically Modern Homo sapiens: Anatomy
Anatomically modern Homo sapiens are characterized by:
A gracile skull
Minimal brow ridges
Rounded cranium, higher forehead

77. Cranial capacity ~1500 cc
Small teeth and jaws
Retracted face with an obvious chin (mental protuberance)
82
Gracile!
These traits do NOT appear as a unit - earliest anatomically
modern remains may have one trait, but not another - do see
regional variations
82
Comparison of the crania of Homo heidelbergensis, Homo
neandertalensis, and Homo sapiens showing important
differences
56
Anatomically Modern Homo sapiens: Culture
UPPER PALEOLITHIC CULTURE
New stone tool technology

78. Blades and microliths
Earliest dates to 300 kya in Africa
See modern behaviour before modern anatomy
Also abundance of bone, antler, shell, and wood
More hafting of tools
More standardized toolkits of highly specialized implements
Use of atlatl (projectile technology!)
Art!
“explosion” in Europe around 40 kya
84
Atlatl – spear throwing technology
Atlatl in Alberta:
https://albertashistoricplaces.wordpress.com/2015/10/07/alberta
s-ancient-darts-and-atlatl-hunting/
85
Art: Cave Paintings

79. After 40 kya
Location
Deep cold caves
Sound
Isolation - darkness
Ritual parallels
Means of achieving altered state (shamans & trances)
Monastic practices
Vision quests
Sympathetic (hunting) magic
Fertility magic
86
Cave Art
Includes spectacular images of animals and abstract forms and,
rarely, humans
Mobiliary & Other Forms of Art
Depictions of Women
Venus figures
Hohlenstein figure
87

80. Mobiliary Art = portable art objects
Clothing & Ornamentation
Body Ornamentation:
Pierced shells, pierced animal teeth, and bone beads were most
likely work as necklaces or attached to clothing
Lice & clothing
“Human body lice reveal the birthdate of fashion”
Stoneking et al. 2003
Genetics of lice
42-70 kya
88
Humans only started wearing clothes as little as 40,000 years
ago, according to a new genetic study which has calculated
when the human body louse evolved - a creature which needs
clothes to lay its eggs on.
According to the research, by Professor Mark Stoneking and
colleagues at the Max Planck Institute for Evolutionary
Anthropology in Leipzig, Germany and published in the latest
issue of the journal Current Biology, humans might have first
worn clothes around 42,000 to 72,000 years ago.
Anthropologists have long wondered when clothes began to
appear. Since fur and fabrics do not fossilize, no evidence has

81. been left, apart from some fabrics more than a few thousand
years old.
The new approach focused on the subtle genetic differences
between the head louse (Pediculus humanus capitis) and the
body louse (P. humanus corporis or P. h. humanus). These
human ectoparasites differ mainly in their habitat on the host:
head lice live in the hair and scalp, while body lice feed on
hairless parts of the body but lay their eggs only in clothes.
"This ecological differentiation probably arose when humans
adopted frequent use of clothing," write the researchers. Thus,
an indirect measure of when our ancestors first wore clothes
would have emerged by figuring out when body lice first
appeared, the researchers concluded.
DNA analysis of the 40 human head lice and body lice sent from
around the world revealed the modern genetic variation in the
parasites.
Assuming that mutations occur at a given rate, Stoneking's team
came to the estimate that "body lice originated not more than
about 72,000 to 42,000 years ago." The date fits with fossil and
archaeological evidence: the only tools that can be definitely
associated with clothing, such as needles, are about 40,000
years old.
The genetic results also indicate greater diversity in African
than non-African lice, suggesting an African origin of human
lice which matches human origins.
What is a Denisovan, and why do they seem to matter?
Named after Denisova Cave in Russia
Some Neandertal fossils, and another Archaic Homo sapiens
more closely related to Neandertals than Anatomically modern
humans

82. Originate in Asia
Contributed to Asian, Melanesian, Aboriginal Australian and
Native American DNA
DNA markers used to track migrations and divergence of
populations
89
Complex stratigraphy at Denisova Cave
Morely, M.W. et al. Nature Scientific Reports volume 9,
Article number: 13785 (2019) |
90
Map showing potential archaic hominin territories and possible
directions of gene flow for the basal mtDNA haplogroups. This
would reflect migratory movement during the post-Toba climate
event in Eurasia, several millennia prior to the repopulating of
the continent 60Kya.
Following successful recovery of ancient DNA and subsequent
mapping of the Denisovan genome, we now know that these
archaic humans had sexual encounters with our ancestors around
45,000 to 50,000 years ago. The confirmation of interbreeding
with Denisovans has followed the realisation that these early
modern human groups mated with our other close relatives the
Neanderthals. Today most non-Africans have around 2–3%

83. Neanderthal genes; East Asians have the highest levels with
about 30% more of these genes than Europeans.
Denisovan genes are not present in all living humans; they are
at their highest levels among the indigenous people of
Australasia, especially those of Papua where they make up
approximately 5% of the modern genome. Southeast Asians and
East Asians are the only other populations with significant
traces of Denisovan genes with a mere 0.2%. The discovery that
Papuans carry such remarkably high levels of this archaic
human DNA has led many scientists to suspect the Denisovans
interbred with ancestors of Australasians somewhere east of
Wallace’s Line – perhaps on Papua itself or a neighbouring
island.
The paper Analysis of Human Sequence Data Reveals Two
Pulses of Archaic Denisovan Admixture by Browning et al.
(2018), published in the academic journal Cell adds a twist to
the tale of these archaic romantic liaisons. Scientists associated
with the University of Washington contrasted samples of DNA
from numerous modern populations against the genome
extrapolated from the samples of Siberian Denisovan DNA. The
result of this exhaustive process was that the team discovered
that East Asians, particularly Chinese and Japanese people,
carry genes from two distinct Denisovan populations.
The data in the new paper makes it clear that the ancestors of
East Asian people carry genes from the same Southern
Denisovan group reflected in the Papuan genome, but also genes
from a more distant relative of theirs which was more closely
related to the individuals inhabiting the Denisova Cave in
Siberia. That the ancestors of East Asians have this unique
signature for two interbreeding events has caused considerable
confusion.
The logical deduction from this new finding is that some of the
modern humans living in Australasia moved up into Southeast
and East Asia after interbreeding with Southern Denisovan had
occurred. Members of this migrant group then encountered a
second population of Northern Denisovans and had sexual

84. relations with them, even if only on a limited scale – suggested
by the low proportion of genes remaining today. Samples of
DNA from the Denisova Cave site had previously revealed one
of the Denisovans, a young girl, carried genes from a modern
human ancestor closely related to modern Papuan. This
discovery confirms that the genetic traffic moved in both
directions.
These migrants from Australasia must have flourished in Asia
because today they have several million descendants. They also
may have absorbed additional human sub-species previously
living in the region as around one-quarter of the ancient DNA
that Browning found didn’t match either Neanderthal or
Denisovan DNA. It could be that various so-called ‘ghost
populations’ mated with early modern humans in Asia, we can’t
be sure until we find fossils containing viable DNA.
90
Relationship Status: It’s Complicated!
91
Stringer 2015
Next Topic
Questions ?
Concerns ?
Problems ?
Final Exam!
92

85. 92
THE LIVING PRIMATES
1
Primatology: Why Study the Living Primates?
1950s = rise of field of primatology
Primatologist: A scientist working directly or indirectly with
nonhuman primates in a variety of settings
Goals of primatology:
Reconstruct early hominin & human adaptations, behaviors by
providing models & analogues
2

86. -so, who studies primate behavior and how do they do it?
-primatologists are scientists who study the evolution, anatomy,
and behavior of non-human primates
-primatologists can be trained as zoologists or biological
anthropologists
-the earliest studies of nonhuman primates was in the 1920s and
1930s but in the 1950s is when we really see primatology as a
discipline take off
-Robert Yerkes, an American psychologist, sent his students
into the field to study gorillas, chimps, and howler monkeys
2
Are we looking through a window or in a mirror?
http://domain.me/emoticons/
One of the issues we face in primatology is that it is hard to be
an objective observer. One must recognize how our own culture
affects interpretation of primate behaviour: How much do we
see because we want to see it? How much are we missing

87. because we aren’t looking for it?
So beware direct comparisons between primates (and between
living primates and our fossil ancestors):
Modern primates provide clues:
Useful as models, not because they represent an earlier stage in
our evolution as a species (THEY DO NOT), but because they
are all we have for even remotely related mammals – there are
no living populations of related hominins.
We must be critical of interpretation and validity of the models
created (Anth209!).
3
What we have been able to show?
Primates are intelligent
Impressive skills re: tool use
Ability to manipulate symbols
Ape-language experiments
Ability to reason analogically
Ability to learn and transmit knowledge across generations
Sense of self
ability to recognize oneself (the “mirror test”)
4
4
What we have been able to show?
Primates are social
Behave differently to differently dominant individuals

88. Recognize & respond to individual personalities
Recognize kin from non-kin
Ally with close kin first in conflicts
5
5
What we also know . . .
No significant relationship between environment & primate
social organization
Primates are all about dietary & behavioral plasticity
Monogamous pairs
Rare in non-human primates
Those monogamous non-human primates typically lack sexual
dimorphism & have social organization where sex roles are
undifferentiated
6
Note violence (including sexual violence) and homosexuality
has been documented but not in all groups. This demonstrates
that these are a) not exclusive to humans and b) have a
biological component.
6

89. Primate Characteristics & Taxonomy
This is a HUGE field of study so our goals at the 101 level are
to understand
What is a primate
What characteristics do all primates share
How primates vary
What we can learn from non-human primates to better
understand ourselves and our evolution (think forces or
“fingers” of evolution…)
7
PRIMATE CHARACTERISTICS: LeGros Clark (1963)
So what is a primate
-identifying single traits that define the primate order isn’t easy
because compared to most mammals, primates have remained
quite generalized (this is likely an advantage for us...)

90. -that is, primates have retained many ancestral or primitive
mammalian traits that many other mammals have lost over time
-because primates are remarkably generalized, they can’t be
defined by one or even a few traits they share in common
-therefore, anthropologists have drawn attention to a group of
characteristics that, when taken together, more or less
characterize the entire primate order
-still, these are a set of general tendencies that aren’t all equally
expressed in all primates
-in addition, while some of these traits are unique to primates,
many others are retained primitive mammalian traits
-so, to get an overall picture of what makes a primate, people
usually focus on their anatomical, life history, and behavioural
traits
-their limbs and locomotion, teeth and diet, senses, brain, and
behaviour is reflective of a common evolutionary history with
adaptations to similar environmental challenges
Increase in brain size relative to body size and increase in the
complexity of the neocortex.
Reduction in both the projection of the face and the reliance on
the sense of smell
Increasing dependence on the sense of sight resulting in the
relocation of the eyes onto the same plane on the front of the
face so the visual field of each eye overlaps, producing depth
perception (stereoscopic vision)
Reduction in the number of teeth
Grasping (prehensile) hands and feet: includes opposable
thumbs and great toes, nails rather than claws, sensitive pads on

91. tips of fingers and toes, and dermal ridges/friction skin on the
digits, soles, palms, and underside of prehensile tails
Important is distinction between homology (similarities because
of shared ancestry) versus analogy (similar adaptation to similar
selective pressures, not shared ancestry)
8
Primate Characteristics
This table considers these characteristics of primates using
principles from evolutionary theory
9
Lemurs and Lorises
Tarsiers, Monkeys, Apes, Humans
10
See Figures 4.3 (p.73) and 4.12 (p.83)
Cladistic taxonomy of the primates.
-instead of simply moving tarsiers into the suborder
Anthropoidea, one scheme places lemurs and lorises in a

92. different suborder, Strepsirhini instead of Prosimii
-in this classification, tarsiers are included with monkeys, apes,
and humans in another suborder, Haplorhini
-in this classification, the conventionally name suborders
Prosimii and Anthropoidea are replaced by Strepsirhini and
Haplorhini
-this designation has not been universally accepted
so, if you see the term strepshirhini, you know we are referring
to lemurs and lorises
-this classification system makes the biological and
evolutionary statement that all the strepsirhine species are more
closely related to one another than they are to any of the
haplorhines
-likewise, all haplorhine species are more closely related to one
another than they are to any of the strepsirhines
What is represented here as the “alternative” classification has
been increasingly accepted as the correct one thanks to insights
from genetics but also following core principles of cladistics
.
10
Suborder: Strepsirhines
11
Lemurs, Lorises, and Galagos
Old World tropics and subtropics
Retain some primitive traits not seen in other primates
Best living analogy to earliest fossil primates

93. -the first group we are going to talk about is the suborder of
Strepsirhines
-the strepsirhine suborder is made up of the lemurs and lorises
-Strepsirhines are found in throughout the Old World Tropics
and subtropics, in places such as Madagascar, India, and Africa
-Lemurs, Lorises, and galagos, the strepsirhines, are the most
primitive of all primates
-remember that by primitive I mean that strepsihrines, taken as
a group, are more similar anatomically to their earlier
mammalian ancestors than are the other primates (tarsiers,
monkeys, apes, and humans)
-in evolutionary terms, the strepsirhines are among the oldest
living primates, making them a good group to study in order to
understand fossil primates
-so, they tend to retain certain ancestral characteristics, such as
more pronounced reliance on olfaction
-they are also the most varied category of primates
-many characteristics distinguish lemurs and lorises from the
haplorines, including eyes place more to the side of the face,
differences in reproductive physiology, and shorter gestation
periods

94. 11
Suborder: Strepsirhines
12
Characteristic traits:
Good sense of smell (olfaction)
Rhinarium – moist nose
Prehensile hands and feet with nails on most digits
Tooth comb
Stereoscopic vision with orbits enclosed by a postorbital bar
Dental formula: 2133/2133
-many of strepsirhines are nocturnal (active in the night), hence:
They have a more developed sense of smell than other primates
(wet, "naked" doglike nose) – referred to as the rhinarium
large eyes
- all have a "grooming claw" on the second toe only; nails on
other digits
- all have a "dental comb" comprised of the four lower incisor
teeth and the lower canines, which are long, narrow, and close
together, for use in grooming fur and gathering food
-the mandible is also unfused, there is a plate of cartilage in
between, this gives the mandible a v-shaped appearance
-all have the post-orbital bar but lack the post-orbital plate, like
haplorhines do
-most have all three premolars so their dental formula is 2133

95. 12
Suborder: Strepsirhines - LEMURS
Lemurs
Madagascar
60 species exist
5 inches to 2 to 3 feet tall
Varied locomotory patterns
Grooming claw, postorbital bar, tooth comb, unfused mandible,
and 2133/2133
Lemurs (only in Malagasy Republic – ex-Madagascar)
Only diurnal prosimian (competition).
-found only on the island of Madagascar, off the coast of Africa
-60 surviving species of lemurs exist today
-they range in size from the small mouse lemur at 5 inches tall
to the Indri at 2 to 3 feet tall

96. -the larger lemurs are diurnal while the smaller, ones are
nocturnal
-many are primarily arboreal, others such as the ring-tailed
lemur are more terrestrial
-have all the classic strepsirhine traits, grooming claw,
postorbital bar, tooth comb, 2133/2133
13
LORISES & GALAGOS
Lorises (Africa, Asia). Nocturnal.
Galago = Bush Baby
Lorises
Resembles the lemur
8 species of lorises
India, Sri Lanka, Southeast Asia, and Africa
Also includes Galagos
Arboreal quadrupedalism

97. Infant parking
Grooming claw, postorbital bar, tooth comb, unfused mandible,
and 2133/2133
-Lorises, which resemble lemurs, are nocturnal, meaning they
are active during the night
-there are at least eight loris species, all of them found in
tropical forest and woodland habitats of India, Sri Lanka,
Southeast Asia, and Africa
-also included in the same category are 6 to 9 galago species,
which are widely distributed throughout most of the forested
woodland savannah areas of sub Saharan Africa
- locomotion in lorises is a slow, cautious, climbing form of
quadrupedalism
-the flexible hip joints permit suspension by hind limbs while
using the hands in feeding
-all galagos, however, are highly agile vertical clingers and
leapers
-both loris and galago females practice infant parking
-this sounds dangerous, but lorises first bathe their young with
saliva that can cause an allergic reaction sufficient to
discourage most predators
-lorises exhibit good grasping and climbing abilities and well -
developed visual apparatus
-finally, all lorises and galagos exhibit the classic strepsirhine
traits, grooming claw, postorbital bar, tooth comb, 2133/2133

98. 14
Suborder: Haplorhines
15
‘Higher primates’
Tarsiers and Anthropoids (Monkeys, Apes, and Humans)
Share many traits not seen in Strepsirhines
The tarsier is a problem here since it shares many traits with
both Strepsirhines and Haplorhines
-Haplorhines, also considered as the “higher primates”, include
the tarsiers, new and old world monkeys, apes, and humans
-obviously since they are in their own suborder, they share
many traits not seen in the strepsirhine order
-however, the tarsier, posses a problem to taxonomists since it
exhibits traits that are typified in both Strepsirhines and
haplorhines
-so, as we go through these next couple of slides, you will
notice that there will be some comments with regards to the
tarsier, who might not show the complete set of traits that
characterize the haplorhines
but, we have to include it somewhere
15

99. Suborder: Haplorhines
16
Differences from Strepsirhines include:
Increase in body size
Larger brain in relation to body size
Reduced reliance on olfaction
Absence of the rhinarium
Greater degree of colour vision
More generalized dentition
-Haplorhines differ from Strepsirhines in a number of ways
-now we’re going to go over some general traits, don’t forget
that the tarsiers will pose a problem for these generalizatio ns
1. in general, haplorhines such as monkeys, apes, and humans
have a larger average body size
2. they have a larger brain, and of course this is in absolute
terms and relative to body weight
3. they have a reduced reliance on olfaction, so this woul d
reduce the size of the snout
4. they do not have a rhinarium like the strepsirhines did
-this means that the skin found covering the face is also in the
nose area
-the nose is not moist
-5. they also have a greater degree of colour vision

100. 6. they also have a fully enclosed eye orbit in comparison to the
postorbital bar in strepsirhines
7. the mandible is fused giving it a more U-shape (not in
Tarsiers)
8. they definitely have a more generalized dentition, and have
lost the dental comb
-teeth wise, the more advanced haplorhines such as the old
world monkeys, apes, and humans have one less premolar in
each quadrant
16
Suborder: Haplorhines - TARSIERS
Tarsiers
5 recognized species
Restricted to Southeast Asia
Nocturnal
Vertical clinging and leaping
Problem species: blend of characteristics

101. Rotate head 180 degrees
Grooming claw, postorbital plate, unfused mandible, and
2133/2133
-alright, now for the trouble species, the reason for trouble in
classifying primates
-there are five recognized tarsier species, all of which are
restricted to southeast asia
-their name refers to the presence of two highly elongated tarsal
bones in their feet
-these long bones give extra leverage for leaping and searching
for prey
-they inhabit a wide range of forest types, from tropical forest
to backyard garden
-tarsiers are nocturnal and used vertical clinging and leaping to
surprise prey (a lot of insects)
-as already mentioned, tarsiers present a complex blend of
characteristics not seen in other primates
-they’re unique in that their enormous eyes, which dominate
much of the face, are immobile within their sockets
-to compensate for this inability to move their eyes, tarsiers,
like owls, can rotate their heads 180 degrees
-each of their eyes is about the size of their brain!
-similar to the Strepsirhines, tarsiers have a 2133 dental formula
and a grooming claw

102. -unlike the Strepsirhines, the Tarsier has a postorbital plate,
meaning that its orbits are completely enclosed by bone
-also, unlike the Strepsirhines, the tarsier does not have a tooth
comb
17
SUBORDER HAPLORHINES
InFraorders: Platyrrhines and Catarrhines
Platyrrhines:New World
Monkeys
Catarrhines:Old World
Monkeys, Apes, Humans
18
Platyrrhines – New World Primates
Marmosets, tamarins
Claws, not nails
Commonly birth twins
Considered to be monogamous
Squirrel monkeys (Saimiri)
Spider monkeys (Ateles)
Capuchins (Cebus)

103. Howler monkey (Alouatta)
Owl monkey (Aotus)
Titi (Callicebus)
Callitrichidae (callitrichids)
Cebidae (cebids)
19
Found in forested areas in southern Mexico and Central and
South America
Common traits:
Diurnal
Flat nosed/faced
Small bodied, some have prehensile tails
-the new world monkeys can be found in a wide range of
arboreal environments throughout most forested areas in
southern Mexico and Central and Southern America
-they exhibit a wide range of size, diet, and ecological
adaptations
-they range in size from the tiny marmosets and tamarins ~12
ounces to the 20 pound howler monkeys
-New world monkeys are almost exclusively arboreal and some
never come to the ground
-like old world monkeys, all except one species are diurnal

104. -one characteristic that distinguishes New and Old world
monkeys is the shape of the nose
-NWM have broad noses with outward-facing nostrils, whereas
old world monkeys have narrower and more downward facing
nostrils
-because of their outward-facing nostrils, NWM have been
placed into the platyrrhini infraorder and old world monkeys in
the catarhini infraorder
-nwm are called platyrrhini, in French, plat, means flat.. so they
have flat noses
-the new world monkeys have been subdivided into other groups
such as families
-new world monkeys are characteristically small bodied, have 3
premolars like all the other nonhuman primates we have
discussed so far, and are arboreal
-some new world monkeys posses a prehensile tail
-this prehensile tail acts like a 3rd appendage
-the tail can also be used to suspend the body from a branch so
the hands and feet can be used to feed
-anatomically, like I said their dental formula is 2133/2133,
they have a postorbital septum, and have no tooth comb or
grooming claw
-you can see how they are similar to tarsiers…
Two families
19

105. CALLITRICHIDS
Callitrichids
Left = Marmosets
Right = Tamarins
20
CEBIDS
Cebids cont.
Squirrel Monkey: Left

106. Howler Monkey: Right Upper
Titi Monkey: Right Lower
21
Catarrhines – Old World Primates
cata = downward, rhini = nose
5 Families:
Cercopithecidae (e.g. baboons, mandrills)
Colobidae (colobus, langurs, proboscis)
Hylobatidae (gibbons)
Pongidae (orangutan, chimpanzee, gorilla)
Hominidae (Homo sapiens sapiens)
22
-except for humans, old world monkeys are the most widely
distributed of all living primates - they’re found throughout
sub-saharan Africa and southern asia and their habitat ranges
from tropical jungles to semiarid desert and even to seasonally
snow-covered areas in northern japan
-most old world monkeys are quadrupedal and primarily
arboreal, but some such as baboons are also adapted to life on
the ground
-because old world monkeys have downward facing nostrils,
they are placed in the catarrhini infraorder
-locomotion in old world monkeys includes arboreal
quadrupedalism, terrestrial quadrupedalism, and semi -
brachiation and arobatic leaping
-old world monkeys have areas of hardened skin on their

107. buttocks called ischial colosities that serve as sitting pads
-anatomically, old world monkeys have a postorbital plate
-they also lack a tooth comb and grooming claws
-however, unlike all the other non-human primates that we’ve
discussed so far, old world monkeys have 2 premolars, so their
dental formula is 2123/2123, the same as apes and humans, the
next group that we will be discussing
-examples of old world monkeys include langurs, leaf monkeys,
macaques, baboons etc.
22
Family: Cercopithecidae
Cheek pouches (store extra food)
Ischial callosities (sitting pads)
Wide distribution in Africa & Asia
Broad adaptations to arboreal, forest floor, terrestrial, open-
country environments
Most extensive adaptation (excepting Homo) to terrestrial
environment
Some cercopithecids extreme sexual dimorphism
Many in multi-female, multi-male societies
23
23

108. CERCOPITHECIDS
8 Genera
Genus:
Cercopithecus (guenons): Upper Right
Erythrocebus (patas monkeys): Bottom
Papio (baboons): Upper Left
24
CERCOPITHECIDS
8 Genera
Genus:
Mandrillus (mandrills): Upper Left & lower right = example of