34 Vertebrates

Chapter 34
Vertebrates
PowerPoint Lectures for

Biology, Seventh Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
• Overview: Half a Billion Years of Backbones
• By the end of the Cambrian period, some 540

million years ago
– An astonishing variety of animals inhabited
Earth’s oceans
• One of these types of animals

– Gave rise to vertebrates, one of the most
successful groups of animals

• The animals called vertebrates
– Get their name from vertebrae, the series of
bones that make up the backbone
Figure 34.1
• There are approximately 52,000 species of
vertebrates
– Which include the largest organisms ever to
live on the Earth

• Concept 34.1: Chordates have a notochord
and a dorsal, hollow nerve cord
• Vertebrates are a subphylum of the phylum
Chordata
• Chordates are bilaterian animals
– That belong to the clade of animals known as
Deuterostomia
• Two groups of invertebrate deuterostomes, the
urochordates and cephalochordates
– Are more closely related to vertebrates than to
invertebrates
• A hypothetical phylogeny of chordates
Chordates
Craniates
Vertebrates
Gnathostomes
Osteichthyans
(sister group to chordates)
(sharks, rays, chimaeras)

Cephalaspidomorphi
Lobe-fins
(frogs, salamanders)
Tetrapods
Cephalochordata
(ray-finned fishes)
Amniotes
Chondrichthyes
crocodiles, birds)
Echinodermata
(turtles, snakes,
Actinopterygii
Urochordata
(coelacanths)
(lampreys)
(lungfishes)
(hagfishes)
(lancelets)
(tunicates)
(mammals)
Mammalia
Amphibia
Actinistia
Reptilia
Myxini
Dipnoi
Milk
Amniotic egg
Legs
Lobed fins
Lungs or lung derivatives
Jaws, mineralized skeleton
Vertebral column
Head
Brain
Notochord
Figure 34.2 Ancestral deuterostome

Derived Characters of Chordates
• All chordates share a set of derived characters
– Although some species possess some of these
traits only during embryonic development
Dorsal,
hollow
nerve cord
Muscle Notochord Brain

segments
Mouth
Anus
Pharyngeal
slits or clefts
Muscular,
post-anal tail
Figure 34.3
Notochord
• The notochord
– Is a longitudinal, flexible rod located between
the digestive tube and the nerve cord
– Provides skeletal support throughout most of
the length of a chordate
• In most vertebrates, a more complex, jointed

skeleton develops
– And the adult retains only remnants of the
embryonic notochord

Dorsal, Hollow Nerve Cord
• The nerve cord of a chordate embryo
– Develops from a plate of ectoderm that rolls
into a tube dorsal to the notochord
– Develops into the central nervous system: the
brain and the spinal cord

Pharyngeal Slits or Clefts
• In most chordates, grooves in the pharynx
called pharyngeal clefts
– Develop into slits that open to the outside of
the body
• These pharyngeal slits
– Function as suspension-feeding structures in
many invertebrate chordates
– Are modified for gas exchange in aquatic
vertebrates
– Develop into parts of the ear, head, and neck
in terrestrial vertebrates
Muscular, Post-Anal Tail
• Chordates have a tail extending posterior to the
anus
– Although in many species it is lost during
embryonic development
• The chordate tail contains skeletal elements

and muscles
– And it provides much of the propelling force in
many aquatic species

Tunicates
• Tunicates, subphylum Urochordata
– Belong to the deepest-branching lineage of
chordates
– Are marine suspension feeders commonly
called sea squirts

• Tunicates most resemble chordates during
their larval stage
– Which may be as brief as a few minutes
Notochord
Dorsal, hollow
nerve cord
Tail
Excurrent
siphon
Muscle
segments
Incurrent
siphon
Intestine
Stomach
Atrium
Pharynx with slits
Figure 34.4c (c) A tunicate larva is a free-swimming but

nonfeeding “tadpole” in which all four
chief characters of chordates are evident.

• As an adult
– A tunicate draws in water through an incurrent
siphon, filtering food particles
Incurrent
siphon
to mouth
Excurrent
siphon
Excurrent
siphon
Atrium
Pharynx
with
numerous
slits Anus
Intestine
Tunic
Esophagus
Stomach
(a) An adult tunicate, or (b) In the adult, prominent

sea squirt, is a sessile pharyngeal slits function
animal (photo is in suspension feeding,
Figure 34.4a, b approximately life-sized). but other chordate
characters are not obvious.

Lancelets
• Lancelets, subphylum Cephalochordata
– Are named for their bladelike shape
Tentacle 2 cm
Mouth
Pharyngeal slits
Atrium
Notochord
Digestive tract
Dorsal, hollow Atriopore

nerve cord Segmental
muscles
Anus
Tail
Figure 34.5
• Lancelets are marine suspension feeders
– That retain the characteristics of the chordate
body plan as adults

Early Chordate Evolution
• The current life history of tunicates
– Probably does not reflect that of the ancestral
chordate

• Gene expression in lancelets
– Holds clues to the evolution of the vertebrate form
BF1
Otx Hox3
Nerve cord of lancelet

embryo
BF1
Otx Hox3
Brain of vertebrate embryo

(shown straightened)
Midbrain
Figure 34.6 Forebrain Hindbrain
• Concept 34.2: Craniates are chordates that
have a head
• The origin of a head
– Opened up a completely new way of feeding
for chordates: active predation
• Craniates share some common characteristics

– A skull, brain, eyes, and other sensory organs

Derived Characters of Craniates
• One feature unique to craniates
– Is the neural crest, a collection of cells that
appears near the dorsal margins of the closing
neural tube in an embryo
Neural Neural
Dorsal edges
crest tube
of neural plate Ectoderm
Ectoderm
Migrating neural
Notochord crest cells
(a) The neural crest consists of (b) Neural crest cells migrate to
bilateral bands of cells near distant sites in the embryo.
the margins of the embryonic
Figure 34.7a, b folds that form the neural tube.
• Neural crest cells
– Give rise to a variety of structures, including
some of the bones and cartilage of the skull
(c) The cells give rise to some

of the anatomical structures
unique to vertebrates, including
some of the bones and cartilage
of the skull.
Figure 34.7c

The Origin of Craniates
• Craniates evolved at least 530 million years
ago
– During the Cambrian explosion

• The most primitive of the fossils
– Are those of the 3-cm-long Haikouella
(a) Haikouella. Discovered in 1999 in

southern China, Haikouella had eyes
and a brain but lacked a skull, a
Figure 34.8a derived trait of craniates.
• In other Cambrian rocks
– Paleontologists have found fossils of even
more advanced chordates, such as
Haikouichthys 5 mm
(b) Haikouichthys. Haikouichthys had a

Figure 34.8b skull and thus is considered a true craniate.
Hagfishes
• The least derived craniate lineage that still
survives
– Is class Myxini, the hagfishes
Slime glands
Figure 34.9

• Hagfishes are jawless marine craniates
– That have a cartilaginous skull and axial rod of
cartilage derived from the notochord
– That lack vertebrae

• Concept 34.3: Vertebrates are craniates that
have a backbone
• During the Cambrian period
– A lineage of craniates evolved into vertebrates

Derived Characters of Vertebrates
• Vertebrates have
– Vertebrae enclosing a spinal cord
– An elaborate skull
– Fin rays, in aquatic forms

Lampreys
• Lampreys, class Cephalaspidomorphi
– Represent the oldest living lineage of
vertebrates
– Have cartilaginous segments surrounding the
notochord and arching partly over the nerve
cord

• Lampreys are jawless vertebrates
– Inhabiting various marine and freshwater
habitats
Figure 34.10

Fossils of Early Vertebrates
• Conodonts were the first vertebrates
– With mineralized skeletal elements in their
mouth and pharynx
Dorsal view
of head
Dental
Figure 34.11 elements

• Armored, jawless vertebrates called
ostracoderms
– Had defensive plates of bone on their skin
Pteraspis
Pharyngolepis
Figure 34.12
Origins of Bone and Teeth
• Mineralization
– Appears to have originated with vertebrate
mouthparts
• The vertebrate endoskeleton

– Became fully mineralized much later

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

Derived Characters of Gnathostomes
• Gnathostomes have jaws
– That evolved from skeletal supports of the
pharyngeal slits Gill slits Cranium
Mouth
Skeletal rods
Figure 34.13
• Other characters common to gnathostomes
include
– Enhanced sensory systems, including the
lateral line system
– An extensively mineralized endoskeleton
– Paired appendages

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

• 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
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

• 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 Pelvic fins
Pectoral fins
paired pectoral and pelvic 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
Figure 34.15a, b their gaping mouth.
• 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

• Most sharks
– Have a streamlined body and are swift
swimmers
– Have acute senses

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

• Fishes breathe by drawing water over four or
five pairs of gills
– Located in chambers covered by a protective
bony flap called the operculum
Adipose fin Dorsal 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
Ray-Finned Fishes
• Class Actinopterygii, the ray-finned fishes
– Includes nearly all the familiar aquatic
osteichthyans
(a) Yellowfin tuna (Thunnus
albacares), a fast-swimming,
schooling fish that is an important
commercial fish worldwide
(b) Clownfish (Amphiprion

ocellaris), a mutualistic
symbiont of sea anemones
(c) Sea horse (Hippocampus

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

• The fins, supported mainly by long, flexible
rays
– Are modified for maneuvering, defense, and
other functions

Lobe-Fins
• The lobe-fins, class Sarcopterygii
– Have muscular and pectoral fins
– Include coelacanths, lungfishes, and tetrapods
Figure 34.18

• 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

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

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
Figure 34.19 limb
skeleton

• Extraordinary fossil discoveries over the past
20 years
– Have allowed paleontologists to reconstruct
the origin of tetrapods
Millions of years ago
420 415 400 385 370 355 340 325 310 295 280 265
Silurian Devonian Carboniferous Permian To
Paleozoic present
Ray-finned fishes
Coelacanths
Lungfishes
Eusthenopteron
Panderichthys
Elginerpeton
Metaxygnathus
Acanthostega
lchthyostega
Hynerpeton
Greerpeton
Amphibians
Amniotes
Figure 34.20
Amphibians
• Class Amphibia
– Is represented by about 4,800 species of
organisms
• Most amphibians
– Have moist skin that complements the lungs in
gas exchange

• Order Urodela
– Includes salamanders, which have tails
(a) Order Urodela. Urodeles

(salamanders) retain their tail as adults.
Figure 34.21a

• 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

• Order Apoda
– Includes caecilians, which are legless and
resemble worms
(c) Order Apoda. Apodans, or caecilians,
are legless, mainly burrowing amphibians.
Figure 34.21c

• Amphibian means “two lives”
– 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.
• 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

• A phylogeny of amniotes
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Lepidosaurs
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Diapsids Synapsids
Reptiles
Ancestral
amniote
Figure 34.23
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

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

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

Reptiles
• The reptile clade includes
– The tuatara, lizards, snakes, turtles,
crocodilians, birds, and the extinct dinosaurs

• Reptiles
– Have scales that create a waterproof barrier
– Lay shelled eggs on land
Figure 34.25
• 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

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

• As parareptiles were dwindling
– The diapsids were diversifying
• The diapsids are composed of two main

lineages
– The lepidosaurs and the archosaurs

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

• 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
Lepidosaurs
• One surviving lineage of lepidosaurs
– Is represented by two species of lizard-like
reptiles called tuatara
Figure 34.27a (a) Tuatara (Sphenodon punctatus)

• 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
(b) Australian thorny devil

Figure 34.27b
lizard (Moloch horridus)
• Snakes are legless lepidosaurs
– That evolved from lizards
(c) Wagler’s pit viper (Tropidolaemus wagleri), a snake

Figure 34.27c

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

• 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)

Alligators and Crocodiles
• Crocodilians
– Belong to an archosaur lineage that dates
back to the late Triassic
Figure 34.27e (e) American alligator (Alligator mississipiensis)

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

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
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

The Origin of Birds
• Birds probably descended from theropods
– A group of small, carnivorous dinosaurs

• 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

Living Birds
• The ratites, order Struthioniformes
– Are all flightless
(a) Emu. This ratite lives in Australia.
Figure 34.30a

• 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.
• 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

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

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

Early Evolution of Mammals
• Mammals evolved from synapsids
– In the late Triassic period

• 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
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)
Dimetrodon Morganucodon
(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.

• Living lineages of mammals originated in the
Jurassic
– But did not undergo a significant adaptive
radiation until after the Cretaceous

Monotremes
• Monotremes
– Are a small group of egg-laying mammals
consisting of echidnas and the platypus
Figure 34.33

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
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

• 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

• In Australia, convergent evolution
– Has resulted in a diversity of marsupials that
resemble eutherians in other parts of the world
Marsupial mammals Eutherian mammals
Plantigale Deer mouse
Marsupial mole Mole
Sugar glider
Flying squirrel
Wombat
Woodchuck
Wolverine
Tasmanian devil
Patagonian cavy
Kangaroo
Figure 34.35

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

• 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
• 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 Teeth consisting of

Proboscidea 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
Dogs, wolves, teeth and molars for
shearing; carnivorous zebras, tapirs, on each foot;
bears, cats, rhinoceroses herbivorous
weasels, otters,
seals, walruses Indian rhinoceros
Coyote
Chiroptera Adapted for flight; broad

Cetartiodactyla Hooves with an
Bats skinfold that extends
Artiodactyls even number
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 Star-nosed
Pacific white- blubber; carnivorous mole
Figure 34.36 sided porpoise

Primates
• The mammalian order Primates include
– Lemurs, tarsiers, monkeys, and apes
• Humans are members of the ape group

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
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
Figure 34.37
– The tarsiers of Southeast Asia
– The anthropoids, which include monkeys and

hominids worldwide

• The oldest known anthropoid fossils, about 45
million years old
– Indicate that tarsiers are more closely related Anthropoids
to anthropoids
0
Old World monkeys
panzees
New World monkeys
Orangutans
Lemurs, lorises, and pottos
Gibbons
Tarsiers
Gorillas
Humans
Chim-
10
20
30
40
50
60 Ancestral primate
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)

• 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 34.39a, b that open to the sides.

• 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.

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

• Concept 34.8: Humans are bipedal hominoids
with a large brain
• Homo sapiens is about 160,000 years old
– Which is very young considering that life has
existed on Earth for at least 3.5 billion years

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
– Shortened jaw

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

• 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
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

• Hominids originated in Africa
– Approximately 6–7 million years ago
• Early hominids
– Had a small brain, but probably walked upright,
exhibiting mosaic evolution

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

directly to Homo sapiens

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

• Some species walked fully erect
– And had human-like hands and teeth
(b) The Laetoli footprints, more than

(a) Lucy, a 3.24-million-year-old skeleton, 3.5 million years old, confirm that
represents the hominid species upright posture evolved quite early
Australopithecus afarensis. in hominid history.
(c) An artist’s reconstruction of what

A. afarensis may have looked like.
Figure 34.42a–c
Bipedalism
• Hominids began to walk long distances on two
legs
– About 1.9 million years ago

Tool Use
• The oldest evidence of tool use—cut marks on
animal bones
– Is 2.5 million years old

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”

• Homo ergaster
– Was the first fully bipedal, large-brained
hominid
– Existed between 1.9 and 1.6 million years
Figure 34.43
• Homo erectus
– Originated in Africa approximately 1.8 million
years ago
– Was the first hominid to leave Africa

Neanderthals
• Neanderthals, Homo neanderthalensis
– Lived in Europe and the Near East from
200,000 to 30,000 years ago
– Were large, thick-browed hominids
– Became extinct a few thousand years after the

arrival of Homo sapiens in Europe

Homo sapiens
• Homo sapiens
– Appeared in Africa at least 160,000 years ago
Figure 34.44
• The oldest fossils of Homo sapiens outside
Africa
– Date back about 50,000 years ago

• 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

34 Vertebrates

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34 Vertebrates

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Chapter 34

PowerPoint Lectures for

Lectures by Chris Romero

• By the end of the Cambrian period, some 540

• One of these types of animals

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(sister group to chordates)

(sharks, rays, chimaeras)

Lungs or lung derivatives

Jaws, mineralized skeleton

Figure 34.2 Ancestral deuterostome

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Muscle Notochord Brain

• In most vertebrates, a more complex, jointed

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• The chordate tail contains skeletal elements

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Pharynx with slits

Figure 34.4c (c) A tunicate larva is a free-swimming but

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(a) An adult tunicate, or (b) In the adult, prominent

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Dorsal, hollow Atriopore

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Nerve cord of lancelet

Brain of vertebrate embryo

• Craniates share some common characteristics

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(c) The cells give rise to some

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(a) Haikouella. Discovered in 1999 in

(b) Haikouichthys. Haikouichthys had a

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– Fin rays, in aquatic forms

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• The vertebrate endoskeleton

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(a) Coccosteus, a placoderm

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– Were closely related to the ancestors of

(b) Climatius, an acanthodian

• The cartilaginous skeleton

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(a) Blacktip reef shark (Carcharhinus melanopterus).

(b) Southern stingray (Dasyatis americana).

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

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– Control their buoyancy with an air sac known

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Cut edge of Urinary Anal fin