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Chapter 34. cont.

Vertebrates

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


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• Concept 34.4: Gnathostomes are vertebrates that have
jaws
• Today, jawless vertebrates
– Are far outnumbered by those with jaws

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Derived Characters of Gnathostomes
• Gnathostomes have jaws Gill slits
Cranium

– That evolved from


skeletal supports of the Mouth
Skeletal rods
pharyngeal slits

Figure 34.13

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• Other characters common to gnathostomes include
– Enhanced sensory systems, including the lateral line
system (sensitive to vibrations)
– An extensively mineralized endoskeleton
– Paired appendages

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Fossil Gnathostomes
• The earliest gnathostomes in the fossil record
– Are an extinct lineage of armored vertebrates called placoderms

(a) Coccosteus, a placoderm

Figure 34.14a

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• Another group of jawed vertebrates called acanthodians
– Radiated during the Devonian period
– Were closely related to the ancestors of osteichthyans

(b) Climatius, an acanthodian

Figure 34.14b
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Chondrichthyans (Sharks, Rays, and Their Relatives)

• Members of class Chondrichthyes


– Have a skeleton that is composed primarily of cartilage

• The cartilaginous skeleton


– Evolved secondarily from an ancestral mineralized
skeleton

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• The largest and most diverse subclass of Chondrichthyes
– Includes the sharks and rays

(a) Blacktip reef shark (Carcharhinus melanopterus).


Fast swimmers with acute senses, sharks have
paired pectoral and pelvic fins. Pelvic fins
Pectoral fins

(b) Southern stingray (Dasyatis americana).


Most rays are flattened bottom-dwellers that
crush molluscs and crustaceans for food. Some
rays cruise in open water and scoop food into
their gaping mouth.
Figure 34.15a, b

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• A second subclass
– Is composed of a few dozen species of ratfishes

(c) Spotted ratfish (Hydrolagus colliei). Ratfishes,


or chimaeras, typically live at depths greater
than 80 m and feed on shrimps, molluscs,
and sea urchins. Some species have a poisonous
spine at the front of their dorsal fin.

Figure 34.15c

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• Most sharks
– Have a streamlined body and are swift swimmers
– Have acute senses (adaptation with carnivorous
lifestyle)
– Eggs are fertilized internally

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Ray-Finned Fishes and Lobe-Fins
• The vast majority of vertebrates
– Belong to a clade of gnathostomes called Osteichthyes

• Nearly all living osteichthyans


– Have a bony endoskeleton

• Aquatic osteichthyans
– Are the vertebrates we informally call fishes
– Control their buoyancy with an air sac known as a swim bladder

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• Fishes breathe by drawing water over four or five pairs of gills
– Located in chambers covered by a protective bony flap called the
operculum Dorsal fin Adipose fin Caudal
Swim bladder (characteristic of fin
Spinal cord trout)
Brain
Nostril

Cut edge of Urinary Anal fin


operculum Gills Anus bladder
Gonad
Heart
Liver Lateral
Stomach Pelvic fin line
Kidney
Figure 34.16
Intestine
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Ray-Finned Fishes
• Class Actinopterygii, the ray-finned fishes
– Includes nearly all the familiar aquatic osteichthyans
(a) Yellowfin tuna (Thunnus
albacares), a fast-swimming,

• The fins, supported schooling fish that is an important


commercial fish worldwide

mainly by long,
flexible rays
(b) Clownfish (Amphiprion
ocellaris), a mutualistic
– Are modified for symbiont of sea anemones

maneuvering,
(d) Fine-spotted moray eel
defense, and other (c) Sea horse (Hippocampus (Gymnothorax dovii), a
ramulosus), unusual in predator that ambushes
functions the animal kingdom in that prey from crevices in its
the male carries the young coral reef habitat
during their embryonic
Figure 34.17a–d development

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Lobe-Fins
• The lobe-fins, class Sarcopterygii
– Have rod-shaped bones surrounded by thick layer muscle in
pectoral and pelvic fins
– Live in brackish waters
– Lobe fins – help them move across log or muddy bottom
– Include coelacanths, lungfishes, and tetrapods

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Figure 34.18

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• Concept 34.5: Tetrapods are gnathostomes that have limbs and
feet
• One of the most significant events in vertebrate history
– Was when the fins of some lobe-fins evolved into the limbs and
feet of tetrapods

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Derived Characters of Tetrapods
• Tetrapods have some specific adaptations
– Four limbs and feet with digits
– Ears for detecting airborne sounds

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The Origin of Tetrapods
• In one lineage of lobe-fins
– The fins became progressively more limb-like while the rest of
the body retained adaptations for aquatic life

Bones
supporting
gills

Tetrapod
limb
Figure 34.19
skeleton

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• Extraordinary
fossil discoveries
over the past 20
years
– Have allowed
paleontologists
to reconstruct
the origin of
tetrapods

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Amphibians

• Class Amphibia
– Is represented by about 4,800 species of organisms

• Most amphibians
– Have moist skin that complements the lungs in gas
exchange

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• Order Urodela
– Includes salamanders, which have tails

(a) Order Urodela. Urodeles


(salamanders) retain their tail as adults.

Figure 34.21a

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• Order Anura
– Includes frogs and toads, which lack tails

(b) Order Anura. Anurans, such as


this poison arrow frog, lack a tail as adults.

Figure 34.21b

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• Order Apoda
– Includes caecilians, which are legless and
resemble worms

(c) Order Apoda. Apodans, or caecilians,


are legless, mainly burrowing amphibians.

Figure 34.21c

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• Amphibian means “both ways of life”
– A reference to the metamorphosis of an aquatic larva
into a terrestrial adult

(b) The tadpole is


an aquatic
herbivore with
a fishlike tail and
internal gills.

(a) The male grasps the female, stimulating her to (c) During metamorphosis, the
release eggs. The eggs are laid and fertilized in gills and tail are resorbed, and
water. They have a jelly coat but lack a shell and walking legs develop.
Figure 34.22a–c would desiccate in air.
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• Concept 34.6: Amniotes are tetrapods that have a terrestrially
adapted egg
• Amniotes are a group of tetrapods
– Whose living members are the reptiles, including birds, and the
mammals

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• A phylogeny of amniotes

Saurischians

Dinosaurs

Lepidosaurs

Archosaurs

Diapsids Synapsids

Reptiles

Ancestral
amniote

Figure 34.23
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Derived Characters of Amniotes
• Amniotes are named for the major derived character of the
clade, the amniotic egg
– Which contains specialized membranes that protect the embryo
– Allowed the embryo to develop on land

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• The extraembryonic membranes
– Have various functions Extraembryonic membranes

Allantois. The allantois is a disposal Chorion. The chorion and the membrane of the
sac for certain metabolic wastes pro- allantois exchange gases between the embryo
duced by the embryo. The membrane and the air. Oxygen and carbon dioxide diffuse
of the allantois also functions with freely across the shell.
the chorion as a respiratory organ.
Yolk sac. The yolk sac contains the
Amnion. The amnion protects yolk, a stockpile of nutrients. Blood
the embryo in a fluid-filled vessels in the yolk sac membrane transport
cavity that cushions against nutrients from the yolk into the embryo.
mechanical shock. Other nutrients are stored in the albumen (“egg white”).

Embryo

Amniotic cavity
with amniotic fluid

Yolk (nutrients)

Albumen
Shell
Figure 34.24
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• Amniotes also have other terrestrial adaptations
– Such as relatively impermeable skin and the ability to
use the rib cage to ventilate the lungs

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Early Amniotes
• Early amniotes
– Appeared in the Carboniferous period
– Included large herbivores and predators

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Reptiles

• The reptile clade includes


– The tuatara, lizards, snakes, turtles, crocodilians, birds, and the
extinct dinosaurs

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• Reptiles
– Have scales that create a waterproof barrier
– Lay shelled eggs on land

Figure 34.25
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• Most reptiles are ectothermic
– Absorbing external heat as the main source of body
heat

• Birds are endothermic


– Capable of keeping the body warm through metabolism

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The Origin and Evolutionary Radiation of Reptiles
• The oldest reptilian fossils
– Date to about 300 million years ago

• The first major group of reptiles to emerge


– Were the parareptiles, which were mostly large, stocky
herbivores

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• As parareptiles were dwindling
– The diapsids were diversifying

• The diapsids are composed of two main lineages


– The lepidosaurs and the archosaurs

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• The dinosaurs
– Diversified into a vast range of shapes and sizes
– Included the long-necked giants called the theropods

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• Traditionally, dinosaurs were considered slow, sluggish
creatures
– But fossil discoveries and research have led to the
conclusion that dinosaurs were agile and fast moving

• Paleontologists have also discovered signs of parental


care among dinosaurs

Figure 34.26
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Lepidosaurs
• One surviving lineage of lepidosaurs
– Is represented by two species of lizard-like reptiles
called tuatara

Figure 34.27a (a) Tuatara (Sphenodon punctatus)

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• The other major living lineage of lepidosaurs
– Are the squamates, the lizards and snakes

• Lizards
– Are the most numerous and diverse reptiles, apart from
birds

Figure 34.27b (b) Australian thorny devil


lizard (Moloch horridus)
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• Snakes are legless lepidosaurs
– That evolved from lizards

(c) Wagler’s pit viper (Tropidolaemus wagleri), a snake


Figure 34.27c

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Archosaurs - Turtles

• Turtles
– Are the most distinctive group of reptiles alive today

• Some turtles have adapted to deserts


– And others live entirely in ponds and rivers

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• All turtles have a boxlike shell
– Made of upper and lower shields that are fused to the
vertebrae, clavicles, and ribs

Figure 34.27d (d) Eastern box turtle (Terrapene carolina carolina)

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Alligators and Crocodiles
• Crocodilians
– Belong to an archosaur lineage that dates back to the
late Triassic

Figure 34.27e (e) American alligator (Alligator mississipiensis)

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Birds
• Birds are archosaurs
– But almost every feature of their reptilian anatomy has
undergone modification in their adaptation to flight

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Derived Characters of Birds

• Many of the characters of birds


– Are adaptations that facilitate flight

• A bird’s most obvious adaptations for flight


– Are its wings and feathers

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Finger 1

(b) Bone structure

Palm
(a) wing
Finger 2

Forearm
Wrist
Finger 3

Vane Shaft

Shaft Barb
Barbule
Hook
Figure 34.28a–c (c) Feather structure

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The Origin of Birds

• Birds probably descended from theropods


– A group of small, carnivorous dinosaurs

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• By 150 million years ago
– Feathered theropods had evolved into birds

• Archaeopteryx
– Remains the oldest bird known
Wing claw
Toothed beak

Airfoil wing with


contour feathers
Long tail with
Figure 34.29 many vertebrae

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Living Birds
• The ratites, order Struthioniformes
– Are all flightless

(a) Emu. This ratite lives in Australia.

Figure 34.30a

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• The demands of flight
– Have rendered the general body form of many flying
birds similar to one another
(b) Mallards. Like many bird species,
the mallard exhibits pronounced color
differences between the sexes.

(c) Laysan albatrosses. Like most birds,


Laysan albatrosses have specific
mating behaviors, such as this
courtship ritual.

(d) Barn swallows. The barn swallow is a member of


the order Passeriformes. Species in this order are
called perching birds because the toes of their feet
can lock around a branch or wire, enabling the bird
Figure 34.30b–d to rest in place for long periods.
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• Foot structure in bird feet
– Shows considerable variation

Perching bird Grasping bird Raptor Swimming bird


(such as a (such as a (such as a (such as a duck)
cardinal) woodpecker) bald eagle)
Figure 34.31

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• Concept 34.7: Mammals are amniotes that have hair and
produce milk
• Mammals, class Mammalia
– Are represented by more than 5,000 species

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Derived Characters of Mammals
• Mammary glands, which produce milk
– Are a distinctively mammalian character

• Hair is another mammalian characteristic


• Mammals generally have a larger brain
– Than other vertebrates of equivalent size

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Early Evolution of Mammals

• Mammals evolved from synapsids


– In the late Triassic period
– Nonmammalian synapsids lacked hair, had
sprawling gait, laid eggs

• The jaw was remodeled during the evolution of mammals from


nonmammalian synapsids
– And two of the bones that formerly made of the jaw joint were
incorporated into the mammalian middle ear

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Jaw joint Jaw joint Key
Dentary
Angular
Squamosal
Articular
Quadrate

Dimetrodon Morganucodon

(a) The lower jaw of Dimetrodon is composed of several fused bones; two small bones, the quadrate
and articular, form part of the jaw joint. In Morganucodon, the lower jaw is reduced to a single bone,
the dentary, and the location of the jaw joint has shifted.

Middle ear Stapes Inner ear Eardrum Middle ear

Inner ear
Eardrum
Stapes
Incus (evolved
Sound Sound
from quadrate)
Malleus (evolved
from articular)

Present-day reptile Present-day mammal


(b) During the evolutionary remodeling of the mammalian skull, the quadrate and articular bones became incorporated
into the middle ear as two of the three bones that transmit sound from the eardrum to the inner ear. The steps in
Figure 34.32a, b this evolutionary remodeling are evident in a succession of fossils.

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• Living lineages of mammals originated in the Jurassic
– But did not undergo a significant adaptive radiation until
after the Cretaceous
– Early Cretaceous (140mya) – 3 major lineages of
mammals had emerged:
• Monotremes (egg-laying mammals)
• Marsupials (mammals with a pouch)
• Eutherians (placental mammals)

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Monotremes
• Monotremes
– Are a small group of egg-laying mammals
consisting of echidnas and the platypus

Figure 34.33

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Marsupials

• Marsupials
– Include opossums, kangaroos, and koalas

• A marsupial is born very early in its development


– And completes its embryonic development while nursing
within a maternal pouch called a marsupium

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(a) A young brushtail possum. The young of
marsupials are born very early in their
development. They finish their growth
while nursing from a nipple (in their
mother’s pouch in most species).

Figure 34.34a

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• In some species of marsupials, such as the bandicoot
– The marsupium opens to the rear of the mother’s body
as opposed to the front, as in other marsupials
(b) Long-nosed bandicoot. Most bandicoots
are diggers and burrowers that eat mainly
insects but also some small vertebrates and
plant material. Their rear-opening pouch helps
protect the young from dirt as the mother digs.
Other marsupials, such as kangaroos, have a
pouch that opens to the front.

Figure 34.34b

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Marsupial mammals Eutherian mammals
Plantigale Deer mouse

• In Australia, convergent evolution Mole


Marsupial mole

– Has resulted in a diversity of


Flying squirrel
marsupials that resemble Sugar glider

eutherians in other parts of the


world Wombat
Woodchuck

Wolverine
Tasmanian devil

Patagonian cavy
Kangaroo

Figure 34.35

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Eutherians (Placental Mammals)
• Compared to marsupials
– Eutherians have a longer period of pregnancy

• Young eutherians
– Complete their embryonic development within a uterus,
joined to the mother by the placenta

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Phylogenetic relationships of mammals
This clade of eutherians evolved All members of this clade, This is the largest eutherian This diverse clade includes terrestrial
in Africa when the continent which underwent an adaptive clade. It includes the rodents, and marine mammals as well as bats,
was isolated from other radiation in South America, which make up the largest the only flying mammals. A growing
landmasses. It includes belong to the order Xenarthra. mammalian order by far, with body of evidence, including Eocene
Earth’s largest living land One species, the nine-banded about 1,770 species. Humans fossils of whales with feet,
animal (the African elephant), armadillo, is found in the belong to the order Primates. supports putting whales in
as well as species that weigh southern United States. the same order (Cetartiodactyla)
less than 10 g. as pigs, cows, and hippos.

Proboscidea Sirenia
Tubulidentata Rodentia Carnivora
Hyracoidea Lagomorpha Cetartiodactyla
Afrosoricida (golden Primates Perissodactyla
moles and tenrecs) Dermoptera Chiroptera
Macroscelidea (flying lemurs) Eulipotyphla
(elephant shrews) Scandentia Pholidota
Monotremata Marsupialia Xenarthra (tree shrews) (pangolins)

Monotremes Marsupials Eutherians

Possible phylogenetic tree of mammals.


All 20 extant orders of mammals are listed
Ancestral mammal at the top of the tree. Boldfaced orders
are explored on the facing page.
Figure 34.36
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The major eutherian orders
ORDERS MAIN ORDERS MAIN
AND EXAMPLES CHARACTERISTICS AND EXAMPLES CHARACTERISTICS

Monotremata Lay eggs; no Marsupialia Embryo completes


Platypuses, nipples; young Kangaroos, development in
echidnas suck milk from opossums, pouch on mother
fur of mother koalas

Echidna Koala

Tubulidentata
Proboscidea Teeth consisting of
Long, muscular Aardvark
Elephants many thin tubes
trunk; thick, cemented together;
loose skin; upper eats ants and termites
incisors elongated
as tusks
African elephant Aardvark

Aquatic; finlike Hyracoidea Short legs; stumpy tail;


Sirenia
forelimbs and Hyraxes herbivorous; complex,
Manatees,
no hind limbs; multichambered
dugongs
herbivorous stomach
Rock hyrax
Manatee

Rodentia Chisel-like, continuously


Xenarthra Reduced teeth or
Squirrels, growing incisors worn
Sloths, no teeth; herbivorous
beavers, rats, down by gnawing;
anteaters, (sloths) or carnivorous
porcupines, herbivorous
armadillos (anteaters,
armadillos) mice
Red squirrel
Tamandua

Lagomorpha Primates Opposable thumbs;


Chisel-like incisors; Lemurs, forward-facing eyes;
Rabbits, hind legs longer than
hares, picas monkeys, well-developed
forelegs and adapted apes, cerebral cortex;
for running and humans omnivorous
jumping
Golden lion
tamarin
Jackrabbit

Sharp, pointed canine Perissodactyla Hooves with an


Carnivora Horses, odd number of toes
teeth and molars for
Dogs, wolves, zebras, tapirs, on each foot;
shearing; carnivorous
bears, cats, rhinoceroses herbivorous
weasels, otters,
seals, walruses Indian rhinoceros
Coyote

Chiroptera
Cetartiodactyla Hooves with an Adapted for flight; broad
Bats
Artiodactyls even number skinfold that extends
Sheep, pigs of toes on each from elongated fingers
cattle, deer, foot; herbivorous to body and legs;
giraffes carnivorous or
Frog-eating bat herbivorous
Bighorn sheep

Aquatic; streamlined Eulipotyphla Diet consists mainly


Cetaceans body; paddle-like “Core insecti- of insects and other
Whales, forelimbs and no vores”: some small invertebrates
dolphins, hind limbs; thick moles, some
porpoises layer of insulating shrews
blubber; carnivorous Star-nosed
Pacific white-
mole
Figure 34.36 sided porpoise

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Primates
• The mammalian order Primates include
– Lemurs, tarsiers, monkeys, and apes

• Humans are members of the ape group

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Derived Characters of Primates
• Most primates
– Have hands and feet adapted for grasping

• Primates also have


– A large brain and short jaws
– Forward-looking eyes close together on the face,
providing depth perception
– Well-developed parental care and complex social
behavior
– A fully opposable thumb
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Living Primates
• There are three main groups of living primates:
– The lemurs of Madagascar and the lorises and pottos of
tropical Africa and southern Asia
– The tarsiers of Southeast Asia
– The anthropoids, which include monkeys
and hominids worldwide

Figure 34.37
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Anthropoids

• The oldest known


anthropoid fossils, about
45 million years old 0

panzees
Orangutans
Old World monkeys
New World monkeys
Lemurs, lorises, and pottos

Gibbons

Humans
Tarsiers

Gorillas

Chim-
– Indicate that tarsiers are 10

more closely related to

Millions of years ago


anthropoids 20

30

40

50

60 Ancestral primate
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Figure 34.38
• The fossil record indicates that monkeys
– First appeared in the New World (South America) during the
Oligocene

• The first monkeys


– Evolved in the Old World (Africa and Asia)

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• New World and Old World monkeys
– Underwent separate adaptive radiations during their many
millions of years of separation

(a) New World monkeys, such as spider (b) Old World monkeys lack a prehensile tail, and their nostrils
monkeys (shown here), squirrel monkeys, and open downward. This group includes macaques (shown here),
capuchins, have a prehensile tail and nostrils mandrills, baboons, and rhesus monkeys.
Figure 34.45 that open to the sides.

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• The other group of anthropoids, the hominoids
– Consists of primates informally called apes
(a) Gibbons, such as this Muller's gibbon, are
found only in southeastern Asia. Their very
long arms and fingers are adaptations for
brachiation.

(b) Orangutans are shy, solitary apes that live in the rain
forests of Sumatra and Borneo. They spend most of
their time in trees; note the foot adapted for grasping (c) Gorillas are the largest apes: some
and the opposable thumb. males are almost 2 m tall and weigh
about 200 kg. Found only in Africa, these
herbivores usually live in groups of up to
about 20 individuals.

(e) Bonobos are closely


related to chimpanzees
but are smaller. They
(d) Chimpanzees live in tropical Africa. They
survive today only in the
feed and sleep in trees but also spend a
African nation of Congo.
great deal of time on the ground. Chimpanzees
Figure 34.40a–e are intelligent, communicative, and social.

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• Hominoids
– Diverged from Old World monkeys about 20–25 million
years ago

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• Concept 34.8: Humans are bipedal hominoids with a large brain
• Homo sapiens is about 200,000 years old
– Which is very young considering that life has existed on Earth for
at least 3.5 billion years

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Derived Characters of Hominids
• A number of characters distinguish humans from other
hominoids
– Upright posture and bipedal locomotion
– Larger brains
– Language capabilities
– Symbolic thought
– The manufacture and use of complex tools
– Reduce jawbones & jaw muscles
– Shorter digestive tract
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The Earliest Humans

• The study of human origins


– Is known as paleoanthropology

• Paleoanthropologists have discovered fossils of approximately


20 species of extinct hominoids
– That are more closely related to humans than to chimpanzees,
known as hominins

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• These species are known as hominids
Paranthropus Homo Homo
robustus neanderthalensis sapiens
0
Paranthropus Homo ?
boisei ergaster
0.5

1.0
Australopithecus
1.5 africanus

2.0
Kenyanthropus
Millions of years ago platyops
2.5
Australopithecus
garhi Homo
3.0 erectus
Australopithecus
anamensis
3.5
Homo Homo
rudolfensis habilis
4.0

4.5
Ardipithecus Australopithecus
ramidus afarensis
5.0

5.5

6.0 Orrorin tugenensis

6.5
Sahelanthropus
tchadensis
Figure 34.41 7.0

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• Hominids originated in Africa
– Approximately 6–7 million years ago

• Early hominids
– Had a small brain, but probably walked upright, exhibiting mosaic
evolution

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• Two common misconceptions of early hominids include
– Thinking of them as chimpanzees
– Imagining human evolution as a ladder leading directly to Homo
sapiens

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Australopiths
• Australopiths are a paraphyletic assemblage of hominids
– That lived between 4 and 2 million years ago

• Some species walked fully erect (was bipedal)


– And had human-like hands and teeth
– However, brain only 1/3 the size of present-day human

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings


(a) Lucy, a 3.24-million-year-old skeleton, (b) The Laetoli footprints, more than
represents the hominid species 3.5 million years old, confirm that
Australopithecus afarensis. upright posture evolved quite early
in hominid history.

(c) An artist’s reconstruction of what


A. afarensis may have looked like.

Figure 34.42a–c
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Bipedalism

• Hominids began to walk long distances on two legs


– About 1.9 million years ago
– Bipedal walking – requires less energy than walking on
all fours

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Tool Use

• The oldest evidence of tool use—cut marks on animal


bones found in Ethiopia
– Is 2.5 million years old

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings


Early Homo
• The earliest fossils that paleoanthropologists place in our genus
Homo
– Are those of the species Homo habilis, ranging in age from about
2.4 to 1.6 million years

• Stone tools have been found with H. habilis


– Giving this species its name, which means “handy man”

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings


• Homo ergaster
– Was the first fully bipedal,
large-brained hominid
– Existed between 1.9 and
1.6 million years

Figure 34.50
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• Homo erectus
– Originated in Africa approximately 1.8 million years ago
– Was the first hominid to leave Africa

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings


Neanderthals
• Neanderthals, Homo neanderthalensis
– Lived in Europe by 350,000 years ago and later spread
to the Near East, central Asia & southern Siberia
– Were large, thick-browed hominids
– Brain larger than that of present-day humans
– Made hunting tools from stone & wood
– Became extinct a few thousand years after the arrival of
Homo sapiens in Europe

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Homo sapiens
• Homo sapiens
– Appeared in Africa at least 160,000 years ago

Figure 34.44
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• The oldest fossils of Homo sapiens outside Africa are
from Middle East
– Date back about 115,000 years ago

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings


• The rapid expansion of our species
– May have been preceded by changes to the brain that made
symbolic thought and other cognitive innovations possible

Figure 34.45
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

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