Campbell: Biology

Twelfth Edition

Chapter 34
The Origin and Evolution of
Vertebrates

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 CONCEPT 34.1: Chordates have a
notochord and a dorsal, hollow nerve
cord
• Chordates (phylum Chordata) are bilaterian animals
belonging to the clade called Deuterostomia
• Chordates comprise all vertebrates and two groups of
invertebrates: urochordates and cephalochordates
• There are more than 60,000 species of vertebrates,
animals with a backbone
• Vertebrates include the largest and heaviest animals to
have ever lived, but the smallest is less than a centimeter

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Figure 34.2 Phylogeny of living
chordates

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 Derived Characters of Chordates
• All chordates share a set of derived characters, but they
may appear only during early development
• Four key characters of chordates
– Notochord
– Dorsal, hollow nerve cord
– Pharyngeal slits or clefts
– Muscular, post-anal tail

Figure 34.3 Chordate

characteristics

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• The notochord is a longitudinal, flexible rod between the
digestive tube and nerve cord
• It provides flexible skeletal support during early
development and in adults that retain it
• Most vertebrate adults develop a complex, jointed skeleton
and retain only remnants of the notochord

• The chordate nerve cord develops from a plate of

ectoderm rolled into a neural tube dorsal to the notochord
• The nerve cord of the embryo develops into the central
nervous system: the brain and the spinal cord

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• All chordate embryos have grooves along the outer
surface of the pharynx called pharyngeal clefts
• In most, these grooves develop into pharyngeal slits that
open into the pharynx
• The slits allow water to pass from the mouth to the outside
of the body, bypassing the digestive tract
• Functions of pharyngeal slits and arches
– Suspension-feeding structures in many invertebrate
chordates
– Modified into gills for gas exchange in non-tetrapod
vertebrates
– Tetrapods do not develop pharyngeal slits, the arches
develop into parts of the ear, head, and neck
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• Chordates have a tail posterior to the anus that contains
skeletal elements and muscles
• It provides propelling force in many aquatic species
• The tail is greatly reduced during embryonic development
in many species

Lancelets
• Lancelets (Cephalochordata) are marine suspension
feeders, named for their bladelike shape
• They swim using a simplified form of the mechanism used
by fishes
• Adults can reach 6 cm in length and retain key
characteristics of the chordate body plan
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Figure 34.4 The lancelet
Branchiostoma, a cephalochordate

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Tunicates
• Tunicates (Urochordata) are more closely related to other
chordates than are lancelets
• Chordate characteristics are most apparent during the
larval stage, which may last only a few minutes
• Larvae use tail muscles and notochord to swim until they
settle on a suitable substrate for metamorphosis
• Adult tunicates, called sea squirts, are sessile

Figure 34.5 A tunicate, a urochordate

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Early Chordate Evolution
• Ancestral chordates likely resembled lancelets, which
retain all key chordate traits as adults
• Lancelets have a slightly swollen tip on the anterior end of
the nerve cord, instead of a full-fledged brain
• The brain develops under control of the same Hox genes
and pattern of expression as vertebrates
• Ancestral chordates likely had genes associated with
vertebrate organs including the heart and thyroid
• These genes are found in tunicates and vertebrates, but
absent from nonchordate invertebrates

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CONCEPT 34.2: Vertebrates are
chordates that have a backbone
• A skeletal system and complex nervous system have
allowed vertebrates efficiency at two essential tasks
– Capturing food
– Evading predators

Derived Characters of Vertebrates

• Living vertebrates have shared derived characters
distinguishing them from other chordates
– Fore example, they have two or more sets of Hox
genes; lancelets and tunicates have only one
• This additional genetic complexity enabled evolution of the
nervous system and skeleton innovations
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 Derived Characters of Vertebrates
• The neural crest—unique
to vertebrates—appears
along the edges of the
closing neural tube
• Neural crest cells disperse
through the embryo and
give rise to many features
including
– Teeth
– Some bones and
cartilage of the skull
– Several types of
neurons Figure 34.7 The neural crest, embryonic
– Sensory capsules source of many unique vertebrates traits
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Hagfishes and Lampreys
• Hagfishes (Myxini) and
lampreys (Petromyzontida)
are the only living
vertebrates lacking jaws
• Members of these groups
also lack a backbone, but
do have rudimentary
vertebrae
• Molecular analysis supports
their placement as a clade
of jawless vertebrates, the
cyclostomes
Figure 34.UN03 In-text figure, Myxini and
Petromyzontida mini-tree
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• Hagfishes have vertebrae cartilaginous skull, and a mouth
contains tooth-like keratin formations
• Adults retain the notochord as a flexible rod of cartilage
• Lampreys have reduced vertebrae, a notochord, and a
cartilaginous skeleton
• Parasitic species clamp their mouths onto host fish,
feeding on blood and tissue

Figure 34.8 A hagfish Figure 34.9 A sea lamprey

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Early Vertebrate Evolution
• 530 million-year-old fossils from the Cambrian explosion
document the transition to vertebrates
• Fossils of Haikouella, 3-cm-long suspension feeders that
resembled lancelets, are the most primitive
• They had a well-formed brain, eyes, and muscular
segments, but lacked a skull and ear organs
• Myllokunmingia is the first chordate to have a head
• Ear and eye capsules indicate it had a skull, but it lacked
vertebrae
• The origin of a head—consisting of a brain, eyes and other
sensory organs, and a skull—enabled more complex
movement and feeding behaviors
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Figure 34.10 Fossil of an early chordate

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• Conodonts were among the earliest vertebrates in the fossil record,
dating to 500 million years ago
• Large eyes were used to locate prey and a set of barbed mineralized
hooks at the anterior end of the mouth were used to capture prey
• Later emerging vertebrates (485–359 million years ago) shared traits
with lampreys and conodonts
• Mineralized bone covered their bodies, serving as armor to protect them
from predators; early vertebrate skeletons were composed of
unmineralized cartilage
• All of these jawless, armored vertebrates became extinct by the end of
the Devonian

Figure 34.12 Jawless armored vertebrates

Figure 34.11 A conodont
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CONCEPT 34.3: Gnathostomes are
vertebrates that have jaws
• Today, jawed vertebrates, or gnathostomes, outnumber
jawless vertebrates
• Gnathostomes include sharks and their relatives, ray-
finned fishes, lobe-finned fishes, amphibians, reptiles
(including birds), and mammals
Derived Characteristics of Gnathostomes
• Gnathostomes (“jaw mouth”) have jaws (hinged structures)
with teeth used to grip and slice food
• Jaws may have evolved by modification of the skeletal
rods supporting the pharyngeal (gill) slits

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• Other characters common to gnathostomes include
– Genome duplication, including four sets of Hox genes
– Enlarged forebrain
– Enhanced senses of smell and vision
– Lateral line system (aquatic species), rows of organs
sensitive to vibrations located along body sides

Figure 34.13 Possible step in the evolution of jawbones

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Fossil Gnathostomes
• Gnathostomes appeared in the fossil record from about
440 million years ago
• Several adaptions improved swimming efficiency and
control for prey capture and predator evasion
– Paired fins and a tail
– Dorsal, ventral, and anal fins stiffened by bony fin rays
– A more efficient gas exchange system in the gills
• Early gnathostomes included
now extinct armored vertebrates
called placoderms
(“plate-skinned”)

Figure 34.14 Fossil of an early gnathostome

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• Placoderms were extinct by 359 million years ago, followed
by acanthodians about 70 million years later
• Acanthodians were other jawed vertebrates that radiated
about 444–359 million years ago

• By 420 million years ago, jawed vertebrates had diverged

into the three lineages that survive today
– Chondrichthyans
– Ray-finned fishes
– Lobe-fins

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Chondrichthyans (Sharks, Rays, and
Their Relatives)
• Chondrichthyans have a skeleton composed primarily of
cartilage, often impregnated with calcium
• There are about 1,000
species of chondrichthyans
– Most belong to the
sharks, skates, and
rays
– A few dozen belong
to the ratfishes
(chimaeras)

Figure 34.15 Chondrichthyans

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• Sharks have a streamlined body and are swift swimmers,
but do not maneuver well
• Oil is stored in the liver to help maintain buoyancy, but
continual swimming is necessary to avoid sinking
• The largest sharks and rays are suspension feeders; most
sharks are carnivores that have several rows of sharp
teeth for tearing flesh
• Acute senses of sight and smell, and the ability to detect
electrical fields aid in prey capture
• The reproductive tract, excretory system, and digestive
tract all empty into the cloaca, a single opening to the
outside of the body
• Today, chondrichthyans are severely threatened by
overfishing
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Ray-Finned Fishes and Lobe-Fins
• The vast majority of vertebrates are osteichthyans, nearly
all of which have a bony endoskeleton
• Osteichthyans include the bony fishes and tetrapods
• Aquatic osteichthyans are the vertebrates we informally
call fishes
• Water is drawn into the mouth and over the gills by muscle
contractions and movement of the operculum, a bony flap
that protects the gills
• An air sac called the swim bladder is filled to maintain
buoyancy

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 Figure 34.17 Ray-finned fishes (Actinopterygii)
• There are over 27,000
species of ray-finned
fishes (Actinopterygii),
including most familiar
osteichthyans
• Ray-finned fishes face
several threats from humans
– Fish are an important
source of protein; fishing
on an industrial scale has
led to many fishery
collapses
– Damming of rivers hampers
fishes’ access to food,
migratory pathways, and
spawning grounds
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Lobe-Fins (Sarcopterygii)
• Pectoral and pelvic fins have rod-shaped bones
surrounded by a thick layer of muscle
• The lobed fins are used to maneuver across the substrate
of aquatic habitats
• Three lineages surviving today: Coelacanths (Actinistia),
Lungfishes (Dipnoi), Tetrapods
• Coelacanths were thought to have become extinct 75
million years ago
• In 1938, the first living coelacanth was caught off the coast
of South Africa and have been found in various locations
near eastern Africa and Indonesia

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• There are six species of lungfishes in three genera, all
found in the Southern Hemisphere
• They arose in the ocean, but today live in stagnant ponds
and swamps
• Gas exchange occurs in water using gills, or by gulping air
into lungs attached to the pharynx

Figure 34.19 A coelacanth (Latimeria)

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CONCEPT 34.4: Tetrapods are
gnathostomes that have limbs
• The fins of a lineage of lobe-fins evolved into the limbs and
feet of tetrapods by 365 million years ago
• Tetrapods diversified greatly following the evolution of
limbs and the colonization of land
Derived Characters of Tetrapods
• Life on land selected for numerous modifications to the
tetrapod (“four feet”) body plan
– Four limbs and feet with digits
– A neck, enabling independent movement of the head
– Fusion of the pelvic girdle to the backbone
– The absence of gills (except some aquatic species)
– Ears for detecting airborne sounds
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The Origin of Tetrapods
• The tetrapod body plan was likely a modification of the
existing body plan of lobe-fins
• Tiktaalik, nicknamed a “fishapod,” is a fossil species with
characteristics of both fish and tetrapods
• Fossils like
Tiktaalik have
enabled
paleontologists to
reconstruct the
evolution of
tetrapod limbs
from fins

Figure 34.20 Discovery of a “fishapod”: Tiktaalik

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Amphibians
• Amphibians (class
Amphibia) are
represented by
about 6,150
species in three
clades
– Salamanders
(Urodela, “tailed
ones”)
– Frogs (Anura, “tail-
less ones”)
– Caecilians (Apoda,
“legless ones”)

Figure 34.22 Amphibians

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 Lifestyle and Ecology of Amphibians
• The term amphibian means “both ways of life” and refers
to life the life stages of many frog species that live first in
water and then on land
• Frog larvae (tadpoles) are aquatic herbivores with gills, a
lateral line system, and a long, finned tail
• During metamorphosis, tadpoles develop terrestrial
features including legs, lungs, and external eardrums
• The gills and the lateral line system disappear and the
digestive system is adapted to a carnivorous diet
• In addition to lungs, most amphibians have moist skin that
functions in gas exchange

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• A rapid, worldwide decline of amphibian populations has
been documented over the past 30 years
• Causes include a disease-causing chytrid fungus, habitat
loss, climate change, and pollution
• Nine species have become extinct over 40 years
• More than 100 others
are considered possibly
extinct

Figure 34.23 The “dual life” of a frog (Rana temporaria)

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CONCEPT 34.5: Amniotes are tetrapods
that have a terrestrially adapted egg
• Amniotes are
tetrapods
whose living members
are the reptiles
(including birds) and
mammals
• Amniotes are named
for the amniotic egg,
which contains four
membranes that
protect the embryo

Figure 34.25 A phylogeny of amniotes

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Derived Characters of Amniotes
• A key adaptation for life on land, the amniotic egg reduced
dependence on water for reproduction
• The four extraembryonic membranes include the amnion,
chorion, yolk sac, and allantois

• The amnion is a fluid-

filled sac that
surrounds, bathes,
and cushions the
embryo
• The other membranes
function in gas
exchange, nutrient
transfer, and waste
storage Figure 34.26 The amniotic egg
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• The amniotic eggs of most reptiles and some mammals
have a shell that slows dehydration in air
• Most mammals have lost the eggshell and develop the
embryo within the mother’s body instead
• Breathing efficiency improved in amniotes due to the use
of a rib cage to ventilate the lungs
• With more efficient lung breathing, amniotes became less
dependent on gas exchange through the skin
• Skin became less permeable, enabling improved water
conservation in the terrestrial habitat

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Reptiles
• The 20,800 living reptile species include tuataras, lizards,
snakes, turtles, crocodilians, and birds
• Most are squamates (lizards and snakes; 10,425 species)
or birds (10,000 species)
• Reptiles share several derived characters
– Scales containing keratin protect skin from desiccation
and abrasion
– Most lay shelled eggs on land; the shell protects the
egg from drying out
– Fertilization occurs internally, before the shell is
secreted

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• Most reptiles are ectothermic, absorbing external heat as the
main source of body heat
• Ectotherms regulate their body temperature through behavioral
adaptations
• Birds are endothermic, capable of maintaining body temperature
through metabolic activity

Figure 34.28 Hatching reptiles

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The Origin and Evolutionary Radiation
of Reptiles
• The earliest reptiles, diapsids that resembled lizards, lived
about 310 million years ago
• Diapsids have a pair of holes on either side of the skull
through which muscles attach to the jaw

• The diapsids are composed of three main lineages:

– Turtles
– Lepidosaurs, including living tuataras, lizards, and
snakes, and extinct mososaurs
– Archosaurs, including living crocodilians, and extinct
pterosaurs and dinosaurs

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• Pterosaurs originated in the late Triassic and were the first
tetrapods to exhibit flapping flight
• On land, the dinosaurs diversified into a vast range of
shapes and sizes
• With the exception of birds, dinosaurs were extinct by the
end of the Cretaceous (66 million years ago)
• The large sudden decline may have resulted from an
asteroid or comet impact
• This hypothesis is contested because declines began
millions of years before the Cretaceous ended

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Turtles
• The 351 known species of turtles are most closely related to
crocodilians and birds
• They have lost the holes in the skull that are characteristic
of most diapsids
• The boxlike shell, made of upper and lower shields fused to
the vertebrae, clavicles, and ribs, was acquired in stages
over millions of years

Figure 34.29 Extant reptiles (other than birds)

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Lepidosaurs
• One surviving lineage of lepidosaurs is represented by a
single lizard-like species called the tuatara
• Living tuataras are restricted to 30 islands off the coast of
New Zealand
• They are threatened by introduced rats, which consume
their eggs

Figure 34.29 Extant reptiles (other than birds) Figure 34.29 Extant reptiles (other than birds)
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• There are about 10,425 species of squamates (lizards and
snakes)
• Snakes are descended from lizards with legs; some have
retained vestigial pelvic and limb bones
• Snakes are carnivorous and have adaptations to aid in the
capture and consumption of prey, including
– Chemical sensors
– Sensitivity to ground vibrations
– Heat-detecting organs
– Tongue flicking (fans odors
toward sensors in mouth)
– Toxic venom
– Loosely articulated jawbones
and elastic skin (for
swallowing prey whole)
Figure 34.29 Extant reptiles (other than birds)
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Crocodilians
• Crocodilians (alligators and crocodiles) belong to a lineage
that dates back to the late Triassic
• Early species were small terrestrial quadrupeds; later
species were larger and aquatic
• The 24 species of living crocodilians are restricted to warm
regions of the globe

Figure 34.29 Extant reptiles (other than birds)

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Birds
• Birds are archosaurs that have evolved extensive
modifications in their adaptation to flight
• There are about 10,000 species of birds worldwide

Derived Characters of Birds

• Birds have many weight-saving adaptations that improve
the efficiency of flight
– No urinary bladder
– Only one ovary (females)
– Small gonads (both sexes)
– Toothless mouths
– Air-filled bones with honeycombed internal structure

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• Feathers made of β-keratin are arranged to form wings into
airfoils
• Large pectoral muscles anchored to a keel on the sternum
power flapping wings
• Wings may be specialized for soaring or speed
• Birds also have color vision, acute eyesight, and fine
muscle control to
support flight
• Flight benefits
hunting and
scavenging, escape
from terrestrial predators,
and long-distance
migration
Figure 34.30 Form fits function: the avian wing and feather
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• Birds belong to a group of bipedal dinosaurs called
theropods
• Feathers evolved before powered flight with possible
functions including insulation, camouflage, or courtship
display
• Several groups include one or more flightless species;
ratites are an order of flightless birds including the ostrich,
rhea, kiwi, cassowary, and emu
• Penguins are flightless
birds that use powerful
pectoral muscles and
flap their flipper-like
wings to “fly” in water

Figure 34.32 An emu (Dromaius novaehollandiae) a flightless bird native

to Australia
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 Figure 34.33 A king penguin
(Aptenodytes patagonicus) “flying” Figure 34.34 Hummingbird feeding while hovering
underwater

Figure 34.35 A specialized beak Figure 34.36 Feet adapted to perching

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CONCEPT 34.6: Mammals are amniotes
that have hair and produce milk
• Mammals are another lineage of amniotes, represented by
about 6,400 known species
• Mammals have many distinctive derived characters
– Mammary glands, which produce milk to feed young
– Hair and a fat layer under the skin for insulation
– Kidneys, which conserve water from wastes
– Endothermy and a high metabolic rate
– Efficient respiratory and circulatory systems
– A large brain-to-body-size ratio
– Extensive parental care
– Teeth modified for shearing, crushing, or grinding

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Early Evolution of Mammals
• Synapsids are amniotes that include mammals
• Two bones that formally made up the jaw joint were
incorporated into the mammalian middle ear
• Large synapsid herbivores
and carnivores arose in
the Permian period
(299–252 million years ago)
• The first true mammals
arose during the Jurassic
period (201–145 million
years ago)

Figure 34.38 The evolution of the mammalian ear bones

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• Small-bodied mammals coexisted with the dinosaurs
during the Jurassic and Cretaceous periods
• The three major lineages of mammals had emerged by
160 million years ago: Monotremes (egg-laying mammals),
Marsupials (mammals with a pouch), and Eutherians
(placental mammals)
• Monotremes are a small group of egg-laying mammals
found only in Australia and New Guinea
• They include four species of
echidnas and one species of
platypus; females lack nipples
and secrete milk from
glands on their bellies;
the babies suck milk from
their fur
Figure 34.39 Short-beaked echidna (Tachyglossus aculeatus), an Australian monotreme
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• Marsupials share many derived characters with
eutherians that are not found among monotremes
– Higher metabolic rates
– Nipples to provide milk Figure 34.40 Marsupials in Australia and
North America
– Birth of live young
– Embryonic development in the uterus
– A placenta for nutrient transfer from
mother to embryo
• Marsupials are born early
and continue to develop
while nursing in a pouch
called the marsupium
• Marsupials existed
worldwide in the Mesozoic
era, but now live only in
Australia and the Americas
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 Eutherians (Placental Mammals)
Compared with marsupials,
eutherians have a more
complex placenta and longer
pregnancies
Young eutherians complete
embryonic development within
a uterus, joined to the mother
by the placenta

Figure 34.42 Exploring Mammalian Diversity

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Primates
• The mammalian order Primates includes lemurs, tarsiers,
monkeys, and apes
• Humans are members of the ape group
Derived Characters of Primates
• Hands and feet adapted for grasping
• Digits with flat nails instead of claws
• Fingers with skin ridges that form fingerprints
• A relatively large brain and short jaws
• Forward-looking eyes close together on the face
• Well-developed parental care
• Complex socialCopyright
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• Many primate characters arose as adaptations to living in
the trees
– Big toes and relatively moveable thumbs enable feet
and hands to grasp branches
– Fully opposable thumbs (monkeys and apes) that can
touch the ventral surface of all four fingers improve
dexterity
– Enhanced depth perception and eye-hand coordination
• There are three main groups of living primates
– Lemurs (Madagascar) and the lorises and bush babies
(tropical Africa and southern Asia)
– Tarsiers (southeastern Asia)
– Anthropoids including monkeys and apes (worldwide)

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Figure 34.43 Verreux’s sifakas
(Propithecus verreauxi), a type of lemur
Figure 34.45 New World monkeys and Old World monkeys
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• Apes diverged from Old World monkeys about 25–30
million years ago
• Apes include gibbons and four genera of great apes:
Pongo (orangutans), Gorrilla (gorillas), Pan (chimpanzees
and bonobos), and Homo (humans)
• Nonhuman apes are found in the Old World tropics

Figure 34.46 Nonhuman apes

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CONCEPT 34.7: Humans are mammals that
have a large brain and bipedal locomotion
• Homo sapiens arose about 200,000 years ago
• Our species is quite young, considering the 3.5-billion-year
history of life on Earth
Derived Characters of Humans
• A number of characters distinguish humans from other
apes
– Upright posture and bipedal locomotion
– Larger brains capable of language, symbolic thought,
and artistic expression
– Production and use of tools
– Reduced jawbones and jaw muscles
– Shorter digestive tract
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Derived Characters of Humans
• Human and chimpanzee genomes are 99% identical, but
differ in expression of 19 regulatory genes
• Regulatory genes turn other genes on and off; such large
effects could account for many differences between
humans and chimpanzees
The Earliest Hominins
• The study of human origins is known as
paleoanthropology
• Hominins are extinct species that are more closely related
to humans than to chimpanzees
• Paleoanthropologists have discovered fossils of about 25
species of extinct hominins

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• Fossils of the oldest known hominin, Sahelanthropus
tchadensis, date to 6.5 million years ago
• Early hominins shared derived traits with humans
– Reduced canine teeth
– Relatively flat faces
– Increasingly upright and bipedal
– Placement of foramen magnum underneath the skull
• Early hominins had small brains—300–400 cm3 in volume,
compared to 1,300 cm3 in modern humans
• Teeth were larger and jaws projected more beyond the upper
face compared to modern humans
• They were also smaller in stature than humans
– For example, Ardipithecus ramidus (4.4 million years old)
was about 1.2 m tall; humans are about 1.7 m tall
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Figure 34.48 “Ardi,” a 4.4-million-year-
old fossil of Ardipithecus ramidus

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The Earliest Hominins
• There are two common misconceptions about human
evolution
1. Early hominins were chimpanzees or evolved from
chimpanzees
Correction: Hominins and chimpanzees are separate
lineages that diverged from a common ancestor
• There are two common misconceptions about human
evolution
2. Human evolution is like a ladder leading directly from
an ancestral ape to Homo sapiens
Correction: There are many branches and some
coexisting species in the human evolutionary tree,
with Homo sapiens as the final survivors
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Australopiths
• Australopiths are a paraphyletic assemblage of hominins
that lived 4–2 million years ago. Included are:
– Australopithecus anamensis
(4.2–3.9 million years ago) is
the earliest known australopith
– A. africanus (3–2.4 million years ago)
walked fully erect and had humanlike
hands and teeth
– A. afarensis (3.2–2.2 million years ago)
was bipedal, had a small brain and
body, and a long lower jaw

Figure 34.49 Evidence that hominins walked

upright 3.5 million years ago

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Bipedalism
• Hominins began to walk long distances on two legs about
1.9 million years ago
• Bipedal walking was energy efficient in the arid
environments inhabited by hominins at the time

Tool Use
• The oldest evidence of tool use, cut marks on animal
bones, is 2.5 million years old
• Fossil evidence indicates tool use may have originated
prior to the evolution of large brains

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Early Homo
• The earliest Homo fossils (2.4–1.6 million years old)
include those of Homo habilis
• H. habilis (“handy man”) was named for the tools found at
its fossil sites
• They had shorter jaws and larger brains than australopiths

• Homo ergaster (1.9–1.5 million years ago) were fully

bipedal, large-brained hominins
• They had long, slender legs with hip joints well adapted for
long-distance walking
• Their teeth were smaller than australopiths, adapted for
eating softer foods

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• Sexual dimorphism (size difference between
sexes) decreased in H. ergaster and later Homo
species
• Reduced sexual dimorphism is associated with
species that undergo more pair-bonding

• Homo erectus was the first hominin to migrate out

of Africa by at least 1.8 million years ago
• They migrated as far as the Indonesian
archipelago
• H. erectus was extinct by 200,000–70,000 years
ago

Figure 34.50 Fossil of Homo ergaster

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 Neanderthals
• Neanderthals, Homo neanderthalensis, lived in Europe
and the Near East from 350,000 to between 40,000 and
28,000 years ago
• Neanderthals were thick-boned with a larger brain than
modern humans
• They buried their dead, and made tools from stone and
wood
• Humans did not descend directly from Neanderthals; fossil
evidence indicates mating occurred between humans and
Neanderthals

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Homo sapiens
• The ancestors of living humans originated in Africa
• The earliest Homo sapiens fossils are specimens from
Ethiopia that are 195,000 and 160,000 years old
• The oldest H. sapiens fossils outside Africa are from the
Middle East, dating to about 180,000 years ago
• Humans spread beyond Africa in one or more waves, first
to Asia, and then Europe and Australia
• Humans first arrived in the New World about 15,000 years
ago

Figure 34.53 A 160,000-year-old fossil of Homo sapiens

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• The rapid expansion of Homo sapiens is credited to
advances in cognition as humans evolved
• They were the first group to show evidence of symbolic
and sophisticated thought
– For example, a 77,000-year-old artistic carving was recently
discovered in a cave in South Africa
– By 30,000 year ago, humans were producing spectacular cave
paintings

Figure 34.55 Art, a human hallmark

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Figure 34.UN11 Summary of key concepts: clade
descriptions

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