Chapter 34

PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero

• 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

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

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 segments Notochord Brain

Mouth Anus Muscular, post-anal tail Pharyngeal slits or clefts

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 Incurrent siphon

Muscle segments 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 Tunic

Anus Intestine Esophagus Stomach

Figure 34.4a, b

(a) An adult tunicate, or sea squirt, is a sessile animal (photo is approximately life-sized).

(b) In the adult, prominent pharyngeal slits function in suspension feeding, but other chordate characters are not obvious.

Lancelets • Lancelets, subphylum Cephalochordata
Are named for their bladelike shape
Tentacle Mouth 2 cm

Pharyngeal slits Atrium Notochord Digestive tract Atriopore Segmental muscles Anus Tail

Dorsal, hollow nerve cord

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




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
Dorsal edges of neural plate Ectoderm Neural crest Ectoderm Neural tube

Notochord (a) The neural crest consists of bilateral bands of cells near the margins of the embryonic Figure 34.7a, b folds that form the neural tube.

Migrating neural crest cells (b) Neural crest cells migrate to distant sites in the embryo.

• 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 skull and thus is considered a true craniate. Figure 34.8b

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

Figure 34.11

Dental elements

• Armored, jawless vertebrates called ostracoderms
Had defensive plates of bone on their skin


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 Gill slits pharyngeal 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 paired pectoral and pelvic fins.

Pectoral fins

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 fin Adipose
Dorsal fin Nostril Spinal cord Brain Swim bladder (characteristic of trout)

Caudal fin

Cut edge of operculum Gills Heart Figure 34.16 Liver


Urinary Anus bladder Lateral line

Anal fin

Kidney Intestine

Stomach Pelvic fin

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

Figure 34.17a–d

(c) Sea horse (Hippocampus ramulosus), unusual in the animal kingdom in that the male carries the young during their embryonic development

(d) Fine-spotted moray eel (Gymnothorax dovii), a predator that ambushes prey from crevices in its 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

Figure 34.19

Tetrapod 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 Devonian Carboniferous Paleozoic Permian Silurian To 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 release eggs. The eggs are laid and fertilized in water. They have a jelly coat but lack a shell and Figure 34.22a–c would desiccate in air.

(c) During metamorphosis, the gills and tail are resorbed, and walking legs develop.

• 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

Figure 34.23
Pa rar ep t ile Tu rtl es Cr o Pte r s


co d

i lia ns os au rs Or nit din hisc os hia au n rs

Reptiles Diapsids


Sa u din risch os ia tha aurs n n b ot ird her s


Bir d Ple s Ich

Ancestral amniote Lepidosaurs Synapsids


ios thy

au rs os Tu ata r Sq u au rs a am

ate s Ma mm als

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


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

Figure 34.27b (b) Australian thorny devil

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 (a) wing Forearm Wrist Vane Shaft Shaft Barb Barbule Hook (c) Feather structure Finger 3 Palm Finger 2

Figure 34.28a–c

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
Toothed beak Wing claw

Airfoil wing with contour feathers Figure 34.29

Long tail with 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 (such as a cardinal) Figure 34.31

Grasping bird (such as a woodpecker)

Raptor (such as a bald eagle)

Swimming bird (such as a duck)

• 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 Eardrum Sound Sound Stapes Inner ear Eardrum Middle ear Inner ear Stapes Incus (evolved from quadrate) Malleus (evolved from articular) Morganucodon Dimetrodon (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 this evolutionary remodeling are evident in a succession of fossils.

Figure 34.32a, b

• 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 Plantigale Eutherian mammals Deer mouse Marsupial mole Mole

Sugar glider Flying squirrel

Wombat Woodchuck

Tasmanian devil



Patagonian cavy

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



Proboscidea Sirenia Tubulidentata Hyracoidea Afrosoricida (golden moles and tenrecs) Macroscelidea (elephant shrews)


Rodentia Lagomorpha Primates Dermoptera (flying lemurs) Scandentia (tree shrews)

Carnivora Cetartiodactyla Perissodactyla Chiroptera Eulipotyphla Pholidota (pangolins)




Ancestral mammal

Figure 34.36

Possible phylogenetic tree of mammals. All 20 extant orders of mammals are listed at the top of the tree. Boldfaced orders are explored on the facing page.

• The major eutherian orders
ORDERS AND EXAMPLES Monotremata Platypuses, echidnas MAIN CHARACTERISTICS Lay eggs; no nipples; young suck milk from fur of mother Echidna Tubulidentata Aardvark ORDERS AND EXAMPLES Marsupialia Kangaroos, opossums, koalas Koala Teeth consisting of many thin tubes cemented together; eats ants and termites Aardvark MAIN CHARACTERISTICS Embryo completes development in pouch on mother

Proboscidea Elephants

Long, muscular trunk; thick, loose skin; upper incisors elongated as tusks African elephant

Sirenia Manatees, dugongs Manatee

Aquatic; finlike forelimbs and no hind limbs; herbivorous

Hyracoidea Hyraxes Rock hyrax

Short legs; stumpy tail; herbivorous; complex, multichambered stomach

Xenarthra Sloths, anteaters, armadillos Tamandua Lagomorpha Rabbits, hares, picas

Reduced teeth or no teeth; herbivorous (sloths) or carnivorous (anteaters, armadillos)

Rodentia Squirrels, beavers, rats, porcupines, mice

Chisel-like, continuously growing incisors worn down by gnawing; herbivorous Red squirrel

Chisel-like incisors; hind legs longer than forelegs and adapted for running and jumping Jackrabbit Sharp, pointed canine teeth and molars for shearing; carnivorous Coyote Hooves with an even number of toes on each foot; herbivorous Bighorn sheep Aquatic; streamlined body; paddle-like forelimbs and no hind limbs; thick layer of insulating blubber; carnivorous

Primates Lemurs, monkeys, apes, humans Golden lion tamarin

Opposable thumbs; forward-facing eyes; well-developed cerebral cortex; omnivorous

Carnivora Dogs, wolves, bears, cats, weasels, otters, seals, walruses

Perissodactyla Horses, zebras, tapirs, rhinoceroses Indian rhinoceros

Hooves with an odd number of toes on each foot; herbivorous

Cetartiodactyla Artiodactyls Sheep, pigs cattle, deer, giraffes

Chiroptera Bats

Frog-eating bat Eulipotyphla “Core insectivores”: some moles, some shrews

Adapted for flight; broad skinfold that extends from elongated fingers to body and legs; carnivorous or herbivorous

Cetaceans Whales, dolphins, porpoises

Diet consists mainly of insects and other small inv ertebrates Star-nosed mole

Figure 34.36

Pacific whitesided 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 to anthropoids
Anthropoids Old World monkeys New World monkeys Lemurs, lorises, and pottos Orangutans Gibbons Tarsiers Gorillas Chimpanzees Humans 0


Millions of years ago





Figure 34.38


Ancestral primate

• 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

Figure 34.39a,

(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. that open to the sides. b

• 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 and the opposable thumb.

(c) Gorillas are the largest apes: some 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.

Figure 34.40a–e

(d) Chimpanzees live in tropical Africa. They feed and sleep in trees but also spend a great deal of time on the ground. Chimpanzees are intelligent, communicative, and social.

(e) Bonobos are closely related to chimpanzees but are smaller. They survive today only in the African nation of Congo.

• 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
0 0.5 1.0 1.5 2.0 Millions of years ago 2.5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 6.5 Orrorin tugenensis Sahelanthropus tchadensis Ardipithecus ramidus Australopithecus afarensis Kenyanthropus platyops Australopithecus garhi Australopithecus anamensis Homo Homo rudolfensis habilis Homo erectus Paranthropus robustus Paranthropus boisei Homo ? ergaster Homo Homo neanderthalensis sapiens

Australopithecus africanus

Figure 34.41


• 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

(a) Lucy, a 3.24-million-year-old skeleton, represents the hominid species Australopithecus afarensis.

(b) The Laetoli footprints, more than 3.5 million years old, confirm that 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

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