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    Primate and Human Evolution

    Chapter 19

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    Olduvai Gorge on the eastern Serengeti Plain, Northern Tanzania

    is often referred to as The Cradle of Mankind

    because of many important hominid discoveries there

    The Cradle of Mankind

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    Who are we? Where did we come from?

    What is the human genealogy?

    These are basic questionsthat we all ask

    Who are we?

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    Many people enjoy tracingtheir own family history as far back as they can,

    similarly paleoanthropologists are discovering,

    based on recent fossil finds

    that the human family tree goes back

    much farther than we thought

    Goes Back Farther

    Than We Thought

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    In fact, a skull found in the African nation ofChad,

    in 2002 and named Sahelanthropus tchadensis

    but nicknamed Tourma,

    which means "hope of life" in the local Goran language,

    has pushed back the origins of humans

    to nearly 7 million years ago

    Another discovery reported in 2006provides strong evidence for

    an ancestor-descendant relationship

    between two early hominid lines, one of which leads to our own human heritage

    Hope of Life

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    So where does this leave us, evolutionarily

    speaking?

    At a very exciting time as we seek to unravel thehistory of our species

    Our understanding of our genealogy

    is presently in flux,

    and each new fossil hominid find

    sheds more light on our ancestry

    Understanding in Flux

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    Apparently human evolution

    is just like that of other groups

    We have followed

    an uncertain evolutionary path

    As new species evolved,

    they filled ecologic niches

    and either gave rise to descendants

    better adapted to the changing environment

    or became extinct

    Our own evolutionary history

    has many dead-end side branches

    Human Evolution

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    We examine the various primate groups,

    in particular the origin and evolution of the

    hominids,

    the group that includes our ancestors

    However, we must point out

    that new discoveries of fossil hominids,

    as well as new techniques for scientific analysis

    are leading to new hypotheses about our ancestry

    New Hypotheses About

    Our Ancestry

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    As recently as 2000,

    the earliest fossil evidence of hominids

    was from 4.4-million-year-old rocks in easternAfrica

    In 2004, discoveries had pushed

    that age back to almost 7 million years Now, new findings in Ethiopia indicate

    a direct link between two early hominid groups

    that were previously thought to be closely related

    Continuing Discoveries

    Change Our Ideas

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    Primatesare difficult to characterize as an

    order

    because they lack the strong specializations

    found in most other mammalian orders

    We can, however, point to several trends

    in their evolution that help define primates

    and are related to their arboreal,

    or tree-dwelling, ancestry

    What Are Primates?

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    These include changes in the skeleton

    and mode of locomotion,

    an increase in brain size,

    a shift toward smaller, fewer,

    and less specialized teeth,

    and the evolution of stereoscopic vision

    and a grasping hand with opposable thumb

    Not all these trends took place in every primate

    group,

    nor did they evolve at the same rate in each group

    Trends in Primates

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    In fact, some primates

    have retained certain primitive features,whereas others show all

    or most of these trends

    Variations

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    The primate order is divided into two suborders

    Theprosimians, or lower primates,

    include the lemurs, lorises, tarsiers, and tree

    shrews,

    while the anthropoids, or higher primates,

    include monkeys, apes, and humans

    Classification of Primates

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    Order Primates:

    Suborder Prosimii: (lower primates) Lemurs,

    lorises, tarsiers, tree shrews

    Suborder Anthropoidea: (Higher primates)

    Monkeys, apes, humans Superfamily Cercopithecoidea: Macaque, baboon,

    proboscis monkey (Old World monkeys)

    Superfamily Ceboidea: Howler, spider, and squirrel

    monkeys (New World monkeys)

    Superfamily Hominoidea: Apes, humans

    Family Pongidae: Chimpanzees, orangutans, gorillas

    Family Hylobatidae: Gibbons, siamangs

    Family Hominidae: Humans

    Classification of Primates

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    Prosimiansare generally small,

    ranging from species the size of a mouse

    up to those as large as a house cat

    They are arboreal, have five digits

    on each hand and foot

    with either claws or nails,

    and are typically omnivorous

    They have large, forwardly directed eyes

    specialized for night vision,

    hence most are nocturnal

    Prosimians

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    Tarsiers areprosimianprimates

    Tarsier

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    Ring-TailedLemur arealsoprosimians

    Ring-Tailed Lemur

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    As their name implies

    pro means "before," and simian means "ape,

    prosimians are the oldest primate lineage,

    and their fossil record extends back to the Paleocene

    During the Eocene prosimians wereabundant, diversified, and widespread

    in North America, Europe, and Asia

    Prosimians

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    Notharctus, a primitive Eocene prosimian

    Eocene Prosimian

    fromNorthAmerica

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    As the continents moved northwardduring the Cenozoic

    and the climate changed from warm tropical

    to cooler midlatitude conditions,the prosimian population decreased

    in both abundance and diversity

    Prosimians Declined in Cooler

    Climate

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    By the Oligocene, hardly any prosimians

    were left in the northern continents

    as the once widespread Eocene populations

    migrated south to the warmer latitudes

    of Africa, Asia, and Southeast Asia

    Presently, prosimians are found

    only in the tropical regions

    of Asia, India, Africa, and Madagascar

    Prosimians Are Tropical

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    Anthropoids evolved from a prosimian lineage

    sometime during the Late Eocene,

    and by the Oligocene

    they were well established

    Anthropoids are divided into three

    superfamilies.

    Anthropoids

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    New World Monkey

    New World Monkeys constitute asuperfamily belonging to thesuborder Anthropoidea

    (anthropoids)

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    Another

    superfamily of theanthropoids:

    the Old Worldmonkeys

    Old Word Monkey

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    The third superfamilyis the great apes,

    which include gorillasand...

    Great Apes

    hi

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    Chimpanzees

    Chimpanzees

    l i f A h id

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    Much of our knowledge about

    the early evolutionary history of anthropoids

    comes from fossils found in the Fayum district,

    a small desert area southwest of Cairo, Egypt

    During the Late Eocene and Oligocene,

    this region of Africa was a lush, tropical rain forest

    that supported a diverse and abundant fauna and

    flora

    Within this forest lived many different

    arboreal anthropoids as well as various prosimians

    Early History of Anthropoids

    Th d f F il S i

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    In fact, several thousand fossil specimens

    representing more than 20 species of primates

    have been recovered from rocks of this region

    One of the earliest anthropoids, and a possible ancestor of the Old World monkeys,

    wasAegyptopithecus,

    a small, fruit-eating, arboreal primate, about 5 kg

    It had monkey characteristics and ape features

    and is the closest link we currently have

    to Old World primates

    Thousands of Fossil Specimens

    O f h E li A h id

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    One of the Earliest Anthropoids

    Skull of

    Aegyptopithecuszeuxis,

    one of the earliestknown anthropoids

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    Anthropoids are divided into three

    superfamilies

    Old World monkeys,

    New World monkeys,

    and hominoids

    Anthropoid Superfamilies

    Cl ifi i f P i

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    Order Primates:

    Suborder Prosimii: (lower primates) Lemurs,

    lorises, tarsiers, tree shrews

    Suborder Anthropoidea: (Higher primates

    Monkeys, apes, humans Superfamily Cercopithecoidea: Macaque, baboon,

    proboscis monkey (Old World monkeys)

    Superfamily Ceboidea: Howler, spider, and squirrel

    monkeys (New World monkeys)

    Superfamily Hominoidea: Apes, humans

    Family Pongidae: Chimpanzees, orangutans, gorillas

    Family Hylobatidae: Gibbons, siamangs

    Family Hominidae: Humans

    Classification of Primates

    Old W ld M k A ib

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    Old World monkeys superfamily Cercopithecoidea

    are characterized by close-set,

    downward-directed nostrils

    like those of apes and humansgrasping hands,

    and a nonprehensile tail

    They include

    the macaque,

    baboon,

    and proboscis monkey

    Old World Monkey Attributes

    Old W d M k

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    Superfamily

    Cercopithecoidea the Old Worldmonkeys

    Old Word Monkey

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    Present-day Old World monkeys

    are distributed in the tropical regions

    of Africa and Asia

    and are thought to have evolved

    from a primitive anthropoid ancestor,such as Aegyptopithecus,

    sometime during the Oligocene

    Old World Monkeys Distribution

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    New World monkeys superfamily Ceboidea

    are found only in Central and South America

    They probably evolved from African monkeys

    that migrated across the widening Atlantic

    sometime during the Early Oligocene,

    and they have continued evolving in isolation

    to this present day

    New World Monkeys

    N W ld M k

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    New World Monkey

    New World Monkeys aremembers of the superfamilyCeboidea

    N C t t

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    No evidence exists of any prosimian

    or other primitive primates

    in Central or South America

    nor of any contact with Old World monkeys

    after the initial immigration from Africa

    New World monkeys are characterized

    by a prehensile tail, flattish face,

    and widely separated nostrils

    and include the howler, spider, and squirrel

    monkeys

    No Contact

    H i id

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    Hominoids

    superfamily Hominoidea

    consist of three families:

    the great apes

    family Pongidae which includes chimpanzees, orangutans, and gorillas

    the lesser apes

    family Hylobatidae

    which are gibbons and siamangs;

    and the hominids

    family Hominidae

    which are humans and their extinct ancestors

    Hominoids

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    The hominoid lineage

    diverged from Old World monkeys

    sometime before the Miocene,

    but exactly when is still being debated

    It is generally accepted, however,

    that hominoids evolved in Africa,

    probably from the ancestral group

    that includedAegyptopithecus

    Hominoid Lineage

    Cli ti Shift

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    Recall that beginning in the Late Eocene

    the northward movement of the continents

    resulted in pronounced climatic shifts

    In Africa, Europe, Asia, and elsewhere,

    a major cooling trend began,

    and the tropical and subtropical rain forests

    slowly began to change to a variety of mixed

    forests

    separated by savannas and open grasslands

    as temperatures and rainfall decreased

    Climatic Shifts

    Apes Adapted

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    As the climate changed,

    the primate populations also changed

    Prosimians and monkeys became rare,

    whereas hominoids diversified

    in the newly forming environmentsand became abundant

    Ape populations became reproductively

    isolatedfrom each other within the various forests,

    leading to adaptive radiation

    and increased diversity among the hominoids

    Apes Adapted

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    During the Miocene,Africa collided with Eurasia,

    producing additional changes in the climate,

    as well as providing opportunitiesfor migration of animals

    between the two landmasses

    Migration of Animals Possible

    Hominoid Relationships

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    Two apelike groups evolved during the

    Miocenethat ultimately gave rise to present-day hominoids

    Although scientists still disagree

    on the early evolutionary relationships among thehominoids,

    fossil evidence and molecular DNA similarities

    between modern hominoid families

    is providing a clearer picture of the evolutionarypathways

    and relationships among the hominoids

    Hominoid Relationships

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    The first group, the dryopithecines,

    evolved in Africa during the Miocene

    and subsequently spread to Eurasia,

    following the collision between the two continents

    The dryopithecines were a varied group of

    hominoids

    in size,

    skeletal features,

    and life-style

    Dryopithecines

    P l

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    The best-known dryopithecine and perhaps

    ancestor of all later hominoids

    isProconsul,

    an ape-like fruit-eating animal

    that led a quadrupedal arboreal existence,

    with limited activity on the ground

    The dryopithecines were very abundantand diverse during the Miocene and Pliocene,

    particularly in Africa

    Proconsul

    Proconsul

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    Proconsul

    Probable appearance of Proconsul, a

    dryopithecine

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    The second group, the sivapithecids,

    evolved in Africa during the Mioceneand then spread throughout Eurasia

    The fossil remains of sivapithecids

    consist mostly of jaws, skulls, and isolated teeth There are few body or limb bones known,

    and thus we know little about their anatomy

    Sivapithecids

    Sivapithecids Ate Harder Foods

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    All sivapithecids had powerful jaws and teethwith thick enamel and flat chewing surfaces,

    suggesting a diet of harder foods such as nuts

    Based on fossil evidence,the sivapithecids were not involved

    in the evolutionary branch leading to humans,

    but were probably the ancestral stock from which present-day orangutans evolved

    In fact, one early genus, Gigantopithecus,

    was a contemporary of earlyHomo in Eastern Asia.

    Sivapithecids Ate Harder Foods

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    Although many pieces are still missing,

    particularly during critical intervals

    in the African hominoid fossil record,

    molecular DNA as well as fossil evidence indicates

    that the dryopithecines, African apes, and hominids

    form a closely related lineage

    The sivapithecids and orangutans

    form a different lineage that did not lead to humans

    Two Lineages

    Hominids

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    The hominids (family Hominidae)

    the primate family that includes present-day humans

    and their extinct ancestors

    have a fossil record extending back

    to almost 7 million years Several features distinguish them from other

    hominoids

    Hominids are bipedal;that is, they have an upright posture,

    which is indicated by several modifications in theirskeleton

    Hominids

    Comparison of Locomotion

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    Comparison betweenquadrupedal and

    bipedal locomotionin gorillas and humans

    Comparison of Locomotion

    In gorillas the ischium

    bone is longand the entire pelvis is

    tilted toward thehorizontal

    Comparison of Locomotion

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    In humans the ischiumbone is much shorter

    and the pelvis is vertical

    Comparison of Locomotion

    Comparison betweenquadrupedal and

    bipedal locomotionin gorillas and humans

    Larger Reorganized Brain

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    In addition, hominids show a trend

    toward a large and internally reorganized brain

    An increase in brain size and organization

    is apparent in comparing the brains of

    a New World Monkey

    Larger Reorganized Brain

    Larger Reorganized Brain

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    In addition, hominids show a trend

    toward a large and internally reorganized brain

    An increase in brain size and organization

    is apparent in comparing the brains of

    Larger Reorganized Brain

    a great ape

    Larger Reorganized Brain

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    In addition, hominids show a trend

    toward a large and internally reorganized brain

    An increase in brain size and organization

    is apparent in comparing the brains of

    Larger Reorganized Brain

    a present-day

    human

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    Other features that distinguish hominids from

    other hominoids includea reduced face

    and reduced canine teeth,

    omnivorous feeding,

    increased manual dexterity,

    and the use of sophisticated tools

    Other Distinguishing Features

    R Cli i Ch

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    Many anthropologists think

    these hominid features evolved in response

    to major climatic changes

    that began during the Miocene

    and continued into the Pliocene

    During this time, vast savannas

    replaced the African tropical rain forests

    where the lower primates

    and Old World monkeys had been so abundant

    Response to Climatic Changes

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    As the savannas and grasslands

    continued to expand,

    the hominids made the transition

    from true forest dwelling

    to life to an environment

    of mixed forests and grasslands

    Mixed Forests and Grasslands

    No Clear Consensus

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    At present, no clear consensus exists

    on the evolutionary history of the hominid lineage

    This is partly because

    to the incomplete fossil record of hominids

    as well as new discoveries,

    and also because some species

    are known only from partial specimens

    or fragments of bone

    Because of this, scientists even disagree

    on the total number of hominid species

    No Clear Consensus

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    A complete discussion

    of all the proposed hominid species

    and the various competing schemes of hominid

    evolution

    is beyond the scope of this course

    However, we will discuss the generally

    accepted taxa

    and present some of the current theories

    of hominid evolution

    Some Current Theories

    Stratigraphic Record

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    The geologic ranges

    for the commonly accepted species of hominids

    Stratigraphic Record

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    Remember that although the fossil record

    of hominid evolution is not complete,

    what does exist is well documented

    Furthermore, it is the interpretation of that

    fossil record

    that precipitates the often vigorous

    and sometimes acrimonious debates

    concerning our evolutionary history

    Debates

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    Discovered in northern Chad's Djurab Desert

    in July, 2002,

    the nearly 7-million-year-old skull

    and dental remains of Sahelanthropus tchadensis

    make it the oldest known hominid yet unearthed

    and very close to the timewhen humans diverged

    from our closest-living relative, the chimpanzee

    Oldest Known Hominid

    Sahelanthropus tchadensis

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    Discovered in

    Chad in 2002

    and dated at

    nearly 7 million

    years,

    this skull is

    presently

    the oldest

    known hominid

    Sahelanthropus tchadensis

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    Currently, most paleoanthropologists accept

    that the human-chimpanzee stock separated

    from gorillas about 8 million years ago

    and humans separated from chimpanzees

    about 5 million years ago

    When Humans and Chimpanzees

    Diverged

    Oldest Hominid

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    Besides being the oldest hominid,

    Sahelanthropus tchadensis shows a mosaic

    of primitive and advanced features

    that has excited and puzzled paleoanthropologists

    The small brain case and most of the teeth(except the canines) are chimplike

    However, the nose, which is fairly flat,

    and the prominent brow ridgesare features only seen, until now,

    in the human genusHomo

    Oldest Hominid

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    Sahelanthropus tchadensis may have been

    bipedal in its walking habits,

    but until bones from its legs and feet are found,

    that supposition remains conjecture

    Leg Bones and Feet Needed

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    The next oldest hominid is Orrorin tugenensis,

    whose fossils have been dated at 6 million years

    and consist of bits of jaw, isolated teeth,

    finger, arm, and partial upper leg bones

    At this time, debate continues

    as to exactly where Orrorin tugenensisfits in the

    hominid lineage

    Next Oldest Hominid

    Ardipithecus ramidus

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    Sometime between 5.8 and 5.2 million years

    ago,

    another hominid was present in eastern Africa

    Ardipithecus ramidus kadabbais older

    than its 4.4 million year old relative

    Ardipithecus ramidus ramidus

    Ardipithecus ramidus kadabbais very similar

    in most features toArdipithecus ramidus ramidus

    but in certain features of its teeth

    is more apelike than its younger relative

    Ardipithecus ramidus

    Stratigraphic Record

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    The geologic age ranges

    for the commonly accepted species of hominids

    Stratigraphic Record

    Habitual Bipedal Walkers

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    Although many paleoanthropologists think

    both Orrorin tugenensisandArdipithecus ramiduskadabba

    were habitual bipedal walkers

    and thus on a direct evolutionary line to humans,

    others are not as impressed with the fossil evidence

    and are reserving judgment

    Until more fossil evidence is found and

    analyzed,any single scheme of hominid evolution presented

    here would be premature

    b u ped W e s

    A t l ith i

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    Australopithecineis a collective term

    for all members of the genusAustralopithecus

    Currently, five species are recognized:

    A. anamensis,

    A. afarensis,

    A. africanus,

    A. robustus,

    and A. boisei

    Australopithecines

    E l ti S h

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    Many paleontologists accept

    the evolutionary scheme in which

    A. anamensis, the oldest known australopithecine,

    is ancestral toA. afarensis,

    who in turn is ancestral toA. africanus

    and the genusHomo,

    as well as the side branch of australopithecines

    represented byA. robustusandA. boisei

    Evolutionary Scheme

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    The oldest known australopithecine

    isAustralopithecus anamensis

    and was discovered at Kanapoi,

    a site near Lake Turkana, Kenya,

    by Meave Leakey

    of the National Museums of Kenya

    and her colleagues

    Oldest Known Australopithecine

    Similar Yet More Primitive

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    A. anamensis,a 4.2-million-year-old bipedalspecies,

    has many features in common

    with its younger relative,A. afarensis,yet is more primitive in other characteristics,

    such as its teeth and skull

    A. anamensis

    is estimated to have been

    between 1.3 and 1.5 m tall

    and weighed between 33 and 50 kg

    New Fossil Discovery

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    A discovery in 2006 of fossils

    ofA. anamensis,from the Middle Awash area in northeastern Ethiopia

    has shed new light on the transition between

    Ardipithecus andAustralopithecus.

    The discovery ofArdipithecus

    in the same region of Africa

    and same times as the earliestAustralopithecus

    provides strong evidence thatArdipithecus evolved intoAustralopithecus

    and links these two genera

    in the evolutionary lineage leading to humans.

    y

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    Australopithecus afarensis,

    who lived 3.93.0 million years ago,

    was fully bipedal

    and exhibited great variability in size and weight

    Members of this species rangedfrom just over 1 m to about 1.5 m tall

    and weighed between 29 and 45 kg

    Australopithecus afarensis

    Lucy

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    A reconstruction of

    Lucys skeleton

    by Owen Lovejoy

    and his students at Kent

    State University, Ohio Lucy is an ~ 3.5-million-

    year-old

    Australopithecusafarensis

    y

    This

    recon-struction

    illustrates how adaptations in

    Lucys hip, leg and foot

    allowed a fully bipedal

    means of locomotion

    Hominid Footprints

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    Preserved in volcanic

    ash at Laetoli, Tanzania Discovered in 1978 by

    Mary Leakey,

    these footprints proved

    hominids

    were bipedal walkers at

    least 3.5 million years

    ago

    The footprints of twoadults and possibly those

    of a child

    are clearly visible in this

    photograph

    p

    Hominid Footprints

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    p

    Most scientists thinkthe footprints

    were made by

    Australopithecusafarensis

    whose fossils are

    found at Laetoli

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    A. afarensishad a brain size of 380450 cubic

    centimeters (cc),

    larger than the 300400 cc

    of a chimpanzee

    but much smaller than that of present-day humans

    (1350 cc average)

    Brain Size ofA. afarensis

    Apelike Features

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    The skull ofA. afarensisretained many apelike

    features,

    including massive brow ridges

    and a forward-jutting jaw,

    but its teeth were intermediate

    between those of apes and humans

    The heavily enameled molarswere probably an adaptation to chewing fruits,

    seeds, and roots

    Apelike Features

    Landscape with A. afarensis

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    p

    Re-creation

    of aPliocenelandscape

    showingmembersof

    Australo-pithecusafarensis

    gatheringand eating

    variousfruits andseeds

    A. africanus Lived 3.02.3 mya

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    A. afarensiswas succeeded byAustralopithecus africanus,

    which lived 3.02.3 million years ago

    The differences between the two species arerelatively minor

    They were both about the same size and

    weight,butA. africanushad a flatter face

    and somewhat larger brain

    A. africanus Lived 3.0 2.3 mya

    Skull ofA. africanus

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    A reconstruction of

    the skullofAustralopithecus

    africanus

    This skull,

    known as that of theTaung Child,

    was discovered byRaymond Dart inSouth Africa in 1924

    and marks thebeginning of modernpaleoanthropology

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    It appears the limbs

    ofA. africanusmay not have beenas well adapted for bipedalism

    as those ofA. afarensis

    Not As Well Adapted for

    Bipedalism

    Robust Species

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    BothA. afarensisandA. africanus

    differ markedly from the so-called robust species

    A. boisei(2.61.0 million years ago)

    andA. robustus(2.01.2 million years ago)

    A. boiseiwas 1.21.4 m tall

    and weighed between 34 and 49 kg

    It had a powerful upper body,

    a distinctive bony crest on the top of its skull,

    a flat face, and the largest molars of any hominids

    Robust Species

    A. robustus Was a Vegetarian

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    A. robustus, in contrast,was somewhat smaller (1.11.3 m tall)

    and lighter (3240 kg)

    It had a flat face, and the crown of its skullhad an elevated bony crest

    that provided additional area

    for the attachment of strong jaw muscles Its broad flat molars indicated

    A. robustuswas a vegetarian

    . obustus Was a Vegeta a

    Australopithecus robustus Skull

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    The skull ofAustralopithecusrobustus

    This species had amassive jaw,

    powerful chewingmuscles,

    and large broad

    flat chewing teeth apparently used

    for grinding upcoarse plant food

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    Most scientists accept the idea

    that the robust australopithecines

    form a separate lineage

    from the other australopithecines

    that went extinct 1 million years ago

    Separate Lineage

    The Human Lineage

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    Homo habilis

    The earliest member of our own genusHomoisHomo habilis,

    who lived 2.5-1.6 million years ago

    Its remains were first found at Olduvai Gorge, Tanzania,

    but it is also known

    from Kenya, Ethiopia, and South Africa

    H. habilisevolved from theA. afarensisandA.africanuslineage

    and coexisted withA. africanus

    for about 200,000 years

    Stratigraphic Record

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    The geologic age ranges

    for the commonly accepted species of hominids

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    H. habilishad a larger brain (700 cc average)

    than its australopithecine ancestors,

    but smaller teeth It was about 1.2-1.3 m tall

    and only weighed 32-37 kg

    Characteristics ofHomo habilis

    Homo Erectus

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    In contrast to the australopithecines andH.

    habilis,

    which are unknown outside Africa,

    Homo erectuswas a widely distributed species,

    having migrated from Africa during the Pleistocene

    Specimens have been found

    not only in Africa

    but also in Europe, India, China ("Peking Man"),

    and Indonesia ("Java Man")

    Homo Erectus

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    H. erectusevolved in Africa 1.8 million years

    ago

    and by 1 million years agowas present in southeastern and eastern Asia,

    where it survived until about 100,000 years ago

    Survived in Asia Until About100,000 Years Ago

    H erectus Differed From Modern

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    AlthoughH. erectusdeveloped regional

    variations in form,

    the species differed from modern humans in several

    ways

    Its brain size of 800-1300 cc,

    though much larger than that ofH. habilis,

    was still less than the average forHomo sapiens

    (1350 cc)

    H. erectus Differed From Modern

    Humans

    Size Similar to Humans

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    H. erectus'sskull was thick-walled,

    its face was massive,

    it had prominent brow ridges,

    and its teeth were slightly larger than those of

    present-day humans

    H. erectuswas comparable to size to modern

    humans,standing between 1.6 and 1.8 m tall

    and weighing between 53 and 63 kg

    S e S a to u a s

    Skull ofHomo erectus

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    A reconstruction ofthe skull ofHomo

    erectus

    a widelydistributed species

    whose remains

    have been found

    in Africa, Europe,

    India, China, and

    Indonesia

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    The archaeological record indicatesthatH. erectuswas a tool maker

    Furthermore, some sites show evidence

    that its members used fire and lived in caves,an advantage for those living

    in more northerly climates

    H. erectusWas a Tool Maker

    Homo erectus Using Tools

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    Recreation of a Pleistocene setting in Europe

    in which members ofHomo erectusare

    using fire and stone tools

    The "Out of Africa" View

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    Debate still surrounds the transition

    fromH. erectusto our own species,Homo sapiensPaleoanthropologists are split into two camps

    On the one side are those who supportthe "out of Africa" view

    According to this camp, early modern humansevolved from a single woman in Africa,

    whose offspring then migrated from Africa, perhaps as recently as 100,000 years ago

    and populated Europe and Asia,

    driving the earlier hominid populations toextinction

    The "Multiregional" View

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    On the other side are those supporting the

    "multiregional" view According to this hypothesis,

    early modern humans did not have an isolatedorigin in Africa,

    but rather established separate populationsthroughout Eurasia

    Occasional contact and interbreeding

    between these populations enabled our species tomaintain its overall cohesiveness,

    while still preserving the regional differences

    in people we see today

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    Regardless of which theory turns out to be

    correct,our species,H. sapiens

    most certainly evolved fromH. erectus

    Homo sapiens Evolved

    FromH. erectus

    Neanderthals

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    Perhaps the most famous of all fossil humans

    are theNeanderthals,who inhabited Europe and the Near East

    from about 200,000 to 30,000 years ago

    Some paleoanthropologists regard theNeanderthals

    as a variety or subspecies of our own species

    (Homo sapiens neanderthalensis),whereas others regard them as a separate species

    (Homo neanderthalensis)

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    In any case, their name comesfrom the first specimens found in 1856

    in the Neander Valley near Dsseldorf, Germany

    Specimens Found in Neander

    Valley

    Neanderthals

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    The most notable difference between

    Neanderthals

    and present-day humans is in the skull Neanderthal skulls were long and low

    with heavy brow ridges, a projecting mouth,

    and a weak, receding chin Their brain was slightly larger on average

    than our own, and somewhat differently shaped

    Neanderthals

    Neanderthal Skull

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    Reconstructed

    Neanderthalskull

    TheNeanderthals

    were characterized

    by prominent heavybrow ridges and weak chin

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    The Neanderthal body was

    more massiveand heavily muscled

    than ours,

    with rather short lower limbs,much like those

    of other cold-adapted people of today

    Cold Adapted

    First Humans in Cold Climates

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    Given the specimens from more than 100 sites,

    we now know Neanderthals

    were not much different from us,

    only more robust

    Europe's Neanderthals were the first humans

    to move into truly cold climates,

    enduring miserably long winters and short

    summers

    as they pushed north into tundra country

    First Humans in Cold Climates

    Burial Ceremony in a Cave

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    Archaeological evidence indicates

    Neanderthals lived in caves

    and participated in ritual burials

    as depicted in this painting of a burial ceremony

    such as occurred approximately 60,000 years agoat Shanidar Cave, Iraq

    Took Care of Their Injured

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    The remains of Neanderthals

    are found chiefly in caves

    and hutlike rock shelters,

    which also contain a variety

    of specialized stone tools and weapons

    Furthermore, archaeological evidence indicates

    that Neanderthals commonly

    took care of their injured and buried their dead,frequently with such grave items

    as tools, food, and perhaps even flowers

    Cro-Magnons

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    About 30,000 years ago,humans closely resembling modern Europeans

    moved into the region inhabited

    by the Neanderthals and completely replaced them Cro-Magnons, the name given to

    the successors of the Neanderthals in France,

    lived from about 35,000 to 10,000 years ago;

    during this period the development of art andtechnology

    far exceeded anything the world had seen before

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    Highly skilled nomadic hunters,

    Cro-Magnons followed the herdsin their seasonal migrations

    They used a variety of specialized tools

    in their hunts, including perhaps the bow and arrow They sought refuge in caves and rock shelters

    and formed living groups of various sizes

    Nomadic Hunters

    Cro-Magnon Camp

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    Pleistocene Cro-Magnon camp in Europe

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    Cro-Magnons were also cave painters

    Using paints made from manganese and ironoxides,

    Cro-Magnon people painted hundreds of scenes

    on the ceilings and walls of cavesin France and Spain,

    where many of them are still preserved today

    Cave Painters

    Painting From a Cave in France

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    Cro-Magnons were very skilled cave painters

    Painting of a horse

    from the cave of Niaux, France

    Cultural Evolution

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    With the appearance of Cro-Magnons,

    human evolution has becomealmost entirely cultural rather than biological

    Humans have spread throughout the worldby devising means to deal with a broad range

    of environmental conditions Since the evolution of the Neanderthals

    about 200,000 years ago,

    humans have gone from a stone culture

    to a technology that has allowed us

    to visit other planets with space probes

    and land astronauts on the Moon

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    It remains to be seen

    how we will use this technology in the future

    and whether we will continue as a species,evolve into another species,

    or become extinct as many groups have before us

    Future

    Summary

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    Theprimatesevolved during the Paleocene Several trends help characterize primate

    and differentiate them from other mammalian

    orders, including a change in overall skeletal structure and

    mode of locomotion

    an increase in brain size

    stereoscopic vision

    and evolution of a grasping hand with opposable

    thumb

    Summary

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    The primates are divided into two suborderstheprosimiansand the anthropoids

    The prosimians are the oldest primate lineage

    and include lemurs, lorises, tarsiers, and treeshrews

    The anthropoids include

    theNewand Old World monkeys,

    apes,

    and hominids, which are humans

    and their extinct ancestors

    Summary

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    The oldest known hominid is Sahelanthropus

    tchadensis, dated at nearly 7 million years

    then two subspecies ofArdipithecusat 5.8 and 4.4 million

    years respectively

    These early hominids were succeeded by the

    australopithecines

    a fully bipedal group that evolved in Africa 4.2 million

    years ago

    Recent discoveries indicateArdipithecus evolved

    intoAustralopithecus

    Summary

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    Currently, five australopithecine species are

    known:Australopithecus anamensis, A. afarensis, A.

    africanus, A. robustus and A. boisei

    The human lineage began

    about 2.5 million years ago in Africawith the evolution ofHomohabilis,

    which survived as a species

    until about 1.6 million years ago

    Homoerectusevolved fromH. habilisabout 1.8 million years ago

    and was the first hominid to migrate out of Africa

    Summary

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    Between 1 and 1.8 million years ago,H.

    erectushad spread to Europe, India, China, and

    Indonesia

    H. erectusused fire, made tools, and livedin caves

    Sometime between 200,000 and 100,000years ago

    Homo sapiensevolved fromH. erectus

    These early humans may be ancestors ofNeanderthals

    Summary

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    Neanderthals were not much different

    from present-day humans,

    only more robust

    and with differently shaped skulls

    They made specialized tools and weapons,apparently took care of their injured,

    and buried their dead

    The Cro-Magnonswere the successorsof the Neanderthals

    and lived from about 35,000-10,000 years ago

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    Summary

    Cro-Magnons were highly skilled nomadic

    hunters,formed living groups of various sizes,

    and were also skilled cave painters

    Modern humans succeeded the Cro-Magnonsabout 10,000 years ago