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13_HumanEvolution.pptx

Human evolution

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Classification of Hominins

Kingdom: Animalia

Subphylum: Vertebrata

Class: Mammalia

Order: Primates

Suborder: Anthropoidea

Superfamily: Hominoidea

Family: Hominidea

Subfamily: Homininae

Tribe: Hominini

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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

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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

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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)

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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

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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.

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First Some Review: How Do Primates Get Around?

Primates exhibit several locomotory patterns:

Vertical Clinging and Leaping

Brachiation

Quadrupedalism

Arboreal vs. terrestrial

Knuckle walking

Bipedalism

Habitual

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-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

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How Do Nonhuman Primates Get Around?

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

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-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

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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

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-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

 

 

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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

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-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

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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?

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-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?

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The Skeletal Evidence of Bipedalism https://youtu.be/Y_o9QFWSTbA

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 helps to absorb shock and adds a propulsive spring

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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)

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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:

-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

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Dating Techniques

For a longer instructional video see: https://youtu.be/gqDHzCj90co

When Did the First Hominins Appear?

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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

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(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

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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

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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

The interconnection of stratigraphic sequences with absolute 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.

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BASIC PRINCIPLES OF STRATIGRAPHY

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Layers are successively deposited, one after another, such that the oldest layers are on the bottom and the youngest on the top

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

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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

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

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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)

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Radiocarbon dating (14C) is the single most important method 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.

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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

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THE CARBON CYCLE & RADIOCARBON HALF-LIFE

Watch: WATCH: Radiometric decay (half-life) video https://www.youtube.com/watch?v=-KYYsknr2Ko&list=PLfpP1vMOJhHWPyPUkHiXlCbnbFQY3IGiY&index=8

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.

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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

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Note:

mya = million years ago

kya = thousand years ago

Who were the first hominins?

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).

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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

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

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

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Sahelanthropus tchadensis

6 – 7 mya

Toros-Menalla, Chad

Partial cranium, no postcranial

Bipedal?

Closest common ancestor to chimps?

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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 built with huge browridges)

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5.8 – 6 mya

Western Kenya

19 specimens of jaw, teeth, finger, arm, and leg bones

Orrorin tugenensis

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-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.

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The genus Ardipithecus

Includes near complete skeleton “Ardi”

Lived in lightly wooded environment

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.

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Ardi’s Foot

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Limb and foot adapted for bipedalism

Elongated toes

Divergent big toe

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Ardi’s Place

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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

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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

Pelvis and femur

Footprints (3.6 mya, Laetoli, Tanzania)

Partially arboreal

Longer arms than legs, curved fingers

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-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

 

-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

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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)

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researchers have divided the entire genus up into gracile and robust forms

 

-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

 

 

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Australopithecus anamensis

4.2 – 3.9 mya

Kenya

-anam: “Lake” in Turkana language

Mosaic of ape and Hominin traits

Most primitive of the genus

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These are the earliest and most primitive members of the genus Australopithecus. They lived between 4.2 and 3.9 mya. Anam 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.

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Australopithecus afarensis

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3.9 – 2.9 mya

Ethiopia and Tanzania

Most well-known early Hominin:

Lucy (AL-288)

Dikika child

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. 

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Australopithecus afarensis

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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

 

-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

 

 

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Australopithecus africanus

3 – 2 mya

South Africa (Taung, Sterkfontein, Makapansgat)

Bipedal

More human-like dentition

Taung child

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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 australopithecine 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.

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Australopithecus sediba

1.95 – 1.78 mya

South Africa

2 partial skeletons

Descended from A. africanus

Derived Homo-like morphology

Ancestral to Homo

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-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

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Australopithecus bahrelghazali

3.5 – 3 mya

Chad

First hominin found outside of East African Rift Valley

Partial fragmentary jaw

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-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

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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?

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-Australopithecus garhi is another of the australopithecines

 

-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

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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

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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 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

 

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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

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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.

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Australopithecus robustus

2.8 – 1 mya

South Africa

Robert Broom

Omnivore

Tools ?

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-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 did 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 what chimps do developed in at least two different lineages of hominins: robust australopithecines and whatever led to us.

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Image showing relationships between Ardipithecus, Australopithecus, and Homo

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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 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?

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Overview of Hominins and Stone Tool Industries

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

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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 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)

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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?

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Early Homo: Variability

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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 (?) 

-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 ? ? ?

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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

Who was Homo erectus?

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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 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

 

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Homo erectus

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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 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.

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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

 

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Homo erectus: Culture

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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?! 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 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)

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Isimila, Tanzania

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Comparisons of estimated cranial capacity (cc) & overall cranial morphology

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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

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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

 

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

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How did Homo sapiens evolve?

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

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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

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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

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

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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

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Archaic Homo sapiens: Culture

Acheulian tool tradition continues

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

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Who were the Neandertals?

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Neandertals & Pop Culture

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Rosny, Quest for Fire, 1911 (1981)

HG Wells, The Grisly Folk, 1927

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

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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

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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

Shorter limbs

Generally more robust skeleton (hyper-robust)

More powerfully muscled

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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

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-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

 

-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.

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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)

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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

 

Does this mean Neandertals climbed trees and jumped on to their prey?

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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 hafting

Varied, regionally diverse tool kits = Binford & Bordes Debate (text)

Compound (hafted) tools

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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% 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

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Neandertal Culture: Speech

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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

 

-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

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Neandertal Culture: Burials

Definitely intentional

Le Moustier

La Ferrassie

Shandiar (IV)

Krapina

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-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 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

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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

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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

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Us or Them?

Differing interpretations of placement in human lineage and intelligence over time

Dumb brutes – flower children – social peers

Neandertal prejudice/racism?

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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

 

-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

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NEANDERTALS AS US

In the 1990’s and into the 2000’s more and more support has led 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 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

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.”

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)

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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 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.

Anatomically Modern Humans

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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

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Anatomically Modern Homo sapiens: Anatomy

Anatomically modern Homo sapiens are characterized by:

A gracile skull

Minimal brow ridges

Rounded cranium, higher forehead

Cranial capacity ~1500 cc

Small teeth and jaws

Retracted face with an obvious chin (mental protuberance)

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Gracile!

These traits do NOT appear as a unit - earliest anatomically modern remains may have one trait, but not another - do see regional variations

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Comparison of the crania of Homo heidelbergensis, Homo neandertalensis, and Homo sapiens showing important differences

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Anatomically Modern Homo sapiens: Culture

UPPER PALEOLITHIC CULTURE

New stone tool technology

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

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Atlatl – spear throwing technology

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Art: Cave Paintings

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

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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

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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

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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 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

Originate in Asia

Contributed to Asian, Melanesian, Aboriginal Australian and Native American DNA

DNA markers used to track migrations and divergence of populations

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Complex stratigraphy at Denisova Cave

Morely, M.W. et al. Nature Scientific Reports volume 9, Article number: 13785 (2019) |

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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% 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 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.

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Relationship Status: It’s Complicated!

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Stringer 2015

Next Topic

Questions ?

Concerns ?

Problems ?

Final Exam!

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