EDGAR FRANCISCO PRIETO PIRAQUIVE

UPN
The Origin and Evolution of Vertebrates
Figure 34.1
 One lineage of vertebrates colonized land
365 million years ago
There are about 52,000 species of
vertebrates, including the largest organisms
ever to live on the Earth (how many beetles
were there?)
Theme: Vertebrates have great disparity, a
wide range of differences within the group
WHY?

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Concept 34.1: Chordates have a
notochord and a dorsal, hollow nerve
cord
Chordates (phylum Chordata) are bilaterian
animals that belong to the clade of animals
known as Deuterostomia
Chordates comprise all vertebrates and two
groups of invertebrates, the urochordates and
cephalochordates

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Figure 34.2
We will explore 11 clades of the phylum Chordata
Echinodermata

ANCESTRAL Cephalochordata

Chordates
DEUTEROSTOME
Urochordata
Notochord
Myxini

Craniates
Common
ancestor of
chordates Petromyzontida

Vertebrates
Head
Chondrichthyes

Gnathostomes
Vertebral column
Actinopterygii

Osteichthyans
Jaws, mineralized skeleton
Actinistia

Lobe-fins
Lungs or lung derivatives
Dipnoi
Lobed fins
Amphibia

Tetrapods
Amniotes
Limbs with digits Reptilia

Amniotic egg Mammalia

Milk

What do you notice that is odd?

Figure 34.3

4 Derived Characters of Chordates

Dorsal,
Muscle hollow
segments nerve cord
Notochord

Mouth
Anus Pharyngeal
Muscular, slits or clefts
post-anal tail
Figure 34.4

Cephalochordata

Cirri

Mouth
Pharyngeal slits
Atrium

Digestive tract
Notochord

1 cm
Atriopore
Dorsal,
Segmental
hollow
muscles
nerve cord • Lancelets (Cephalochordata)
Anus are named for their bladelike
Tail shape
• They are marine suspension
feeders that retain
characteristics of the
chordate body plan as adults
Figure 34.5

Urochordata

Incurrent
Notochord Water flow
siphon
to mouth
Dorsal, hollow Excurrent
nerve cord siphon
Tail Excurrent
siphon Excurrent
siphon Atrium
Incurrent
Muscle Pharynx
siphon
segments with
Intestine Anus numerous
slits
Stomach Intestine
Tunic
Atrium Esophagus
Pharynx with slits Stomach
(a) Tunicate larva (b) Adult tunicate (c) Adult tunicate

• Tunicates (Urochordata) are more closely related to other

chordates than are lancelets
• When attacked, tunicates, or “sea squirts,” shoot water
through their excurrent siphon
Early Chordate Evolution
Ancestral chordates may have resembled
lancelets
The same Hox genes that organize the
vertebrate brain are expressed in the
lancelet’s simple nerve cord tip
Genome sequencing suggests that
Genes associated with the heart and thyroid
are common to all chordates
Genes associated with transmission of nerve
impulses are unique to vertebrates

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Figure 34.8

5 mm

Segmented
muscles

Pharyngeal slits
Figure 34.9

Hagfishes
Slime glands
Hagfishes
The most basal group of craniates is Myxini,
the hagfishes
Hagfishes have a cartilaginous skull and axial
rod of cartilage derived from the notochord,
but lack jaws and vertebrae
They have a small brain, eyes, ears, and tooth-
like formations
Hagfishes are marine; most are bottom-
dwelling scavengers

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Figure 34.10

Lampreys
Lampreys
 Lampreys (Petromyzontida) represent the oldest
living lineage of vertebrates
 They are jawless vertebrates that feed by
clamping their mouth onto a live fish, suck blood
 They inhabit various marine and freshwater
habitats
 They have cartilaginous segments surrounding
the notochord and arching partly over the
nerve cord

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Figure 34.11

Conodonts were the first vertebrates with mineralized

skeletal elements

Dental elements
(within
head)
Concept 34.4: Gnathostomes are
vertebrates that have jaws
Today, jawed vertebrates, or gnathostomes,
outnumber jawless vertebrates
Gnathostomes include sharks and their
relatives, ray-finned fishes, lobe-finned fishes,
amphibians, reptiles (including birds), and
mammals

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Figure 34.13

Gill slits Cranium

Mouth
Skeletal rods
Figure 34.14 Fossil of an early gnathostome.

0.5 m
Figure 34.15

Dorsal fins
Chondrichthyans
(Sharks, Rays, and Their
Relatives)
Pectoral
fins Pelvic fins
(a) Blacktip reef shark (Carcharhinus melanopterus)

(b) Southern stingray (Dasyatis americana)

(c) Spotted ratfish (Hydrolagus colliei)

Chondrichthyans (Sharks, Rays, and Their
Relatives)
Chondrichthyans (Chondrichthyes) have a
skeleton composed primarily of cartilage
The largest and most diverse group of
chondrichthyans includes the sharks, rays,
and skates

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  The reproductive tract, excretory system, and
digestive tract empty into a common cloaca

 Shark eggs are fertilized internally but embryos

can develop in different ways
 Oviparous: Eggs hatch outside the mother’s body
 Ovoviviparous: The embryo develops within the
uterus and is nourished by the egg yolk
 Viviparous: The embryo develops within the uterus
and is nourished through a yolk sac placenta from
the mother’s blood

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Ray-Finned Fishes and Lobe-Fins
The vast majority of vertebrates belong to a
clade of gnathostomes called Osteichthyes
Osteichthyans include the bony fish and
tetrapods
Aquatic osteichthyans are the vertebrates we
informally call fishes

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Figure 34.16

Swim Dorsal fin

bladder Caudal
Spinal cord Adipose fin fin
Brain

Nostril

Cut Anal fin

edge of
operculum Liver
Anus
Gonad Lateral
Gills
Stomach line
Kidney Pelvic
Intestine fin Urinary
Heart bladder
Ray-Finned Fishes
Actinopterygii, the ray-finned fishes, include
nearly all the familiar aquatic osteichthyans
Ray-finned fishes originated during the
Silurian period (444 to 416 million years ago)
The fins, supported mainly by long, flexible
rays, are modified for maneuvering,
defense, and other functions

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Figure 34.17

Yellowfin tuna (Thunnus albacares)

Red
lionfish
(Pterois
volitans)

Common
sea horse
(Hippocampus
ramulosus)

Fine-spotted moray eel

(Gymnothorax dovii)
Figure 34.18

Lobe-Fins

5 cm

Lower Scaly
Dorsal
jaw covering
spine

• The lobe-fins (Sarcopterygii) have muscular pelvic and

pectoral fins
• Lobe-fins also originated in the Silurian period
Figure 34.19
Concept 34.5: Tetrapods are
gnathostomes that have limbs
 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
A neck, which allows separate movement of
the head
Fusion of the pelvic girdle to the backbone
The absence of gills (except some aquatic
species)
Ears for detecting airborne sounds

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Figure 34.20

Fish Tetrapod
Characters Characters
Scales Neck
Fins Ribs
Gills and Fin skeleton
lungs Flat skull
Eyes on top
of skull
Shoulder bones
Ribs
Neck Scales
Head
Eyes on top of skull
Humerus
Flat Ulna
skull Elbow “Wrist”
Radius
Fin Fin skeleton
Figure 34.21

Lungfishes

Eusthenopteron

Panderichthys

Tiktaalik

Acanthostega

Tulerpeton
Limbs
with digits

Amphibians Key to
limb bones
Ulna
Amniotes Radius
Humerus
Silurian PALEOZOIC
Devonian Carboniferous Permian
415 400 385 370 355 340 325 310 295 280 265 0
Time (millions of years ago)
Figure 34.22

Amphibians
• Amphibians (class
Amphibia) are
represented by about (a) Order Urodela (salamanders)
6,150 species
• Order Urodela (b) Order
includes salamanders, Anura
(frogs)
which have tails

(c) Order Apoda

(caecilians)
Figure 34.23
(a) Tadpole

(b) During metamorphosis

(c) Mating adults

  Amphibian means “both ways of life,” referring to the
metamorphosis of an aquatic larva into a terrestrial adult
 Most amphibians have moist skin that complements the
lungs in gas exchange

 Amphibian populations have been declining in recent

decades…why are they so vulnerable?

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 Fertilization is external in most species, and
the eggs require a moist environment
In some species, males or females care for
the eggs on their back, in their mouth, or in
their stomach

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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 mammals

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Figure 34.25

Parareptiles

Turtles

Crocodilians

Archosaurs
Reptiles

Pterosaurs

Ornithischian

Dinosaurs
dinosaurs

Saurischians
Diapsids

Saurischian dinosaurs
other than birds
Birds

ANCESTRAL Plesiosaurs
AMNIOTE
Ichthyosaurs
Lepidosaurs

Tuataras

Squamates
Synapsids

Mammals
Figure 34.26

Extraembryonic membranes

Allantois Chorion
Amnion
Yolk sac

Embryo

Amniotic cavity
with amniotic Yolk
fluid (nutrients)

Shell Albumen
Reptiles
 The reptile clade includes
the tuataras, lizards, snakes,
turtles, crocodilians, birds,
and some extinct groups
 Reptiles have scales that
create a waterproof barrier
 Most reptiles lay shelled
eggs on land

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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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Figure 34.29
(a) Tuatara
(Sphenodon
punctatus)

(b) Australian
thorny devil
lizard (Moloch (c) Wagler’s pit viper
horridus) (Tropidolaemus wagleri)

(e) American alligator (Alligator mississippiensis)

(d) Eastern box turtle

(Terrapene carolina
carolina)
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 characters of birds are adaptations
that facilitate flight
The major adaptation is wings with keratin
feathers
Other adaptations include lack of a urinary
bladder, females with only one ovary, small
gonads, and loss of teeth

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Figure 34.30

Finger 1

(b) Bone structure

Palm
(a) Wing Finger 2
Forearm Finger 3
Wrist
Shaft
Shaft
Barb
Vane Barbule
Hook
(c) Feather structure
Figure 34.31
Toothed beak Wing claw

Airfoil wing
with contour Long tail with
feathers many vertebrae

• The demands of flight have rendered the general

body form of many flying birds similar to one another
Figure 34.33

Behavior and morphology has adapted to fulfill distinct niches

Figure 34.34
Figure 34.35
Figure 34.36
Concept 34.7: Mammals are amniotes
that have hair and produce milk
• Mammals, class Mammalia, are represented by more than 5,300 species

• Derived characters of mammals:

 Mammary glands, which produce milk

 Hair

 A high metabolic rate, due to endothermy

 A larger brain than other vertebrates of equivalent size

 Differentiated teeth

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Figure 34.37
Biarmosuchus,
a synapsid
Key
Temporal Articular
fenestra Quadrate
Jaw joint Dentary
Squamosal
(a) Articular and quadrate bones in the jaw

Middle ear
Inner Eardrum Middle ear
Eardrum Stapes ear

Inner ear
Stapes
Incus (quadrate)
Sound Sound

Malleus (articular)

Present-day reptile Present-day mammal

(b) Articular and quadrate bones in the middle ear
Figure 34.38

Monotremes

• Monotremes are a small group of egg-laying

mammals consisting of echidnas and the
platypus
Marsupials
 Marsupials include opossums, kangaroos, and
koalas
 The embryo develops within a placenta in the
mother’s uterus
 A marsupial is born very early in its development
 It completes its embryonic development while
nursing in a maternal pouch called a
marsupium

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Figure 34.39

(a) A young brushtail possum

(b) Long-nosed bandicoot

 Marsupial Eutherian
mammals mammals
Plantigale Deer mouse

Marsupial mole Mole

Figure 34.40
Convergent
evolution of Sugar glider Flying
squirrel
marsupials
and
eutherians
(placental Wombat
Woodchuck
mammals).

Wolverine
Tasmanian devil

Patagonian cavy
Kangaroo
Figure 34.41a

(5 species)
Monotremes Marsupials
Monotremata

(324 species)
ANCESTRAL Marsupialia
MAMMAL

Proboscidea
Sirenia
Tubulidentata

(5,010 species)
Eutherians
Hyracoidea
Afrosoricida
Macroscelidea

Xenarthra

Rodentia
Lagomorpha
Primates
Dermoptera
Scandentia

Carnivora
Cetartiodactyla
Perissodactyla
Chiroptera
Eulipotyphia
Pholidota
Figure 34.41b
Orders and Examples Main Characteristics Orders and Examples Main Characteristics
Monotremata Lay eggs; no nipples; Marsupialia Completes embryonic
Platypuses, young suck milk from Kangaroos, development in pouch
echidnas fur of mother opossums, on mother’s body
koalas
Echidna Koala
Proboscidea Long, muscular trunk; Tubulidentata Teeth consisting of
Elephants thick, loose skin; upper Aardvarks many thin tubes
incisors elongated cemented together;
as tusks eats ants and termites
African elephant Aardvark
Sirenia Aquatic; finlike fore- Hyracoidea Short legs; stumpy
Manatees, limbs and no hind Hyraxes tail; herbivorous;
dugongs limbs; herbivorous complex, multi-
Manatee Rock hyrax chambered stomach

Xenarthra Reduced teeth or no Rodentia Chisel-like, continuously

Sloths, teeth; herbivorous Squirrels, growing incisors worn
anteaters, (sloths) or carnivorous beavers, rats, down by gnawing;
armadillos (anteaters, armadillos) porcupines, herbivorous
Tamandua mice Red squirrel
Lagomorpha Chisel-like incisors; Primates Opposable thumbs;
Rabbits, hares, hind legs longer than Lemurs, monkeys, forward-facing eyes;
picas forelegs and adapted chimpanzees, well-developed cerebral
for running and jumping; gorillas, cortex; omnivorous
herbivorous humans Golden lion
Jackrabbit tamarin
Carnivora Sharp, pointed canine Perissodactyla Hooves with an odd
Dogs, wolves, teeth and molars for Horses, zebras, number of toes on
bears, cats, shearing; carnivorous tapirs, each foot; herbivorous
weasels, otters, rhinoceroses
seals, walruses Coyote Indian rhinoceros
Cetartiodactyla Hooves with an even Chiroptera Adapted for flight;
broad skinfold that
Artiodactyls number of toes on each Bats extends from elongated
Sheep, pigs, foot; herbivorous fingers to body and
cattle, deer, legs; carnivorous or
Frog-eating bat herbivorous
giraffes Bighorn sheep
Cetaceans Aquatic; streamlined body; Eulipotyphla Eat mainly insects
Whales, paddle-like fore-limbs and “Core and other small
dolphins, no hind limbs; thick layer insectivores”: invertebrates
porpoises Pacific white- of insulating blubber; some moles, Star-nosed
sided porpoise carnivorous some shrews mole
 Derived Characters of Primates
Hands, feet for grasping
Flat nails
A large brain and short jaws
Forward-looking eyes close together on the
face, providing depth perception
Complex social behavior and parental care
A fully opposable thumb (in monkeys and
apes)

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 Living Primates
 There are three main groups of living primates
 Lemurs, lorises, and pottos
 Tarsiers
 Anthropoids (monkeys and apes)

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Figure 34.42
Figure 34.43

Lemurs, lorises,
and bush babies

ANCESTRAL Tarsiers
PRIMATE

Anthropoids
New World monkeys

Old World monkeys

Gibbons

Orangutans

Gorillas

Chimpanzees
and bonobos
Humans

60 50 40 30 20 10 0
Time (millions of years ago)
Figure 34.44

(a) New World monkey: (b) Old World monkey: macaque

spider monkey with prehensile tail
Figure 34.45

(a) Gibbon
(b) Orangutan

(c) Gorilla

(d) Chimpanzees

(e) Bonobos
Concept 34.8: Humans are mammals that
have a large brain and bipedal
locomotion
The species 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 Humans
A number of characters distinguish humans
from other apes
Upright posture and bipedal locomotion
Larger brains capable of language,
symbolic thought, artistic expression, the
manufacture and use of complex tools
Reduced jawbones and jaw muscles
Shorter digestive tract

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 The human and chimpanzee genomes are
99% identical
How can we be this close, yet so different?

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Figure 34.46

Paranthropus Homo Homo Homo

robustus ergaster neanderthalensis sapiens
0 ?
Paranthropus
0.5 boisei

1.0

1.5 Australopithecus
africanus
2.0
Millions of years ago

Kenyanthropus
2.5 platyops
Australopithecus
Australo- garhi Homo erectus
3.0 pithecus
anamensis
3.5
Homo
Homo rudolfensis
4.0 habilis

4.5 • Hominins originated in

Australopithecus
afarensis
5.0
Ardipithecus ramidus
Africa about 6–7 million
5.5
years ago
6.0 Orrorin tugensis

6.5
Sahelanthropus
• Early hominins show
7.0
tchadensis
evidence of small brains
and increasing bipedalism
Figure 34.47: Ardi, 4.4 million years old
 Misconception: Early hominins were
chimpanzees
Correction: Hominins and chimpanzees
shared a common ancestor
Misconception: Human evolution is like a
ladder leading directly to Homo sapiens
Correction: Hominin evolution included
many branches or coexisting species,
though only humans survive today

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Australopiths
 Australopiths are a paraphyletic assemblage of
hominins living between 4 and 2 million years
ago
 Some species, such as Australopithecus afarensis
walked fully erect

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Figure 34.48 Evidence that hominins walked upright 3.5 million years ago.

(a) The Laetoli footprints (b) Artist’s reconstruction of A. afarensis

 Homo erectus originated in Africa by 1.8
million years ago
It was the first hominin to leave Africa

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Neanderthals
Neanderthals, Homo neanderthalensis, lived
in Europe and the Near East from 350,000 to
28,000 years ago
They were thick-boned with a larger brain,
they buried their dead, and they made
hunting tools
Debate is ongoing about the extent to
which genetic material was exchanged
between neanderthals and Homo sapiens

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Figure 34.50
EXPERIMENT
Hypothesis: Neanderthals gave rise to European humans.
Expected Chimpanzees
phylogeny:
Neanderthals
Living Europeans
Other living humans

RESULTS
Chimpanzees

Neanderthal 1
Neanderthal 2
European and other
living humans
Homo Sapiens
Homo sapiens appeared in Africa by
195,000 years ago
All living humans are descended from
these African ancestors

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Figure 34.51
 The oldest fossils of Homo sapiens outside
Africa date back about 115,000 years and
are from the Middle East
Humans first arrived in the New World
sometime before 15,000 years ago
In 2004, 18,000-year-old fossils were found in
Indonesia, and a new small hominin was
named: Homo floresiensis

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  Homo sapiens were the first group to show
evidence of symbolic and sophisticated thought
 In 2002, a 77,000-year-old artistic carving was
found in South Africa

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Figure 34.UN10
Clade Description
Cephalochordata Basal chordates; marine suspension feeders that
nerve cord; pharyngeal slits; post-anal tail (lancelets) exhibit four key derived characters of chordates
Chordates: notochord; dorsal, hollow

Urochordata Marine suspension feeders; larvae display the

Gnathostomes: hinged jaws, four sets of Hox genes

(tunicates) derived traits of chordates
Craniates: two sets of Hox

Myxini Jawless marine organisms; have head that includes

(hagfishes and a skull and brain, eyes, and other sensory organs
relatives)
genes, neural crest
Vertebrates: Dix genes duplication,

Petromyzontida Jawless vertebrates; typically feed by attaching to a

(lampreys) live fish and ingesting its blood
backbone of vertebrae

Chondrichthyes Aquatic gnathostomes; have cartilaginous skeleton,

Osteichthyans: bony skeleton

(sharks, rays, a derived trait formed by the reduction of an

skates, ratfishes) ancestral mineralized skeleton
Actinopterygii Aquatic gnathostomes; have bony skeleton and
Lobe-fins: muscular fins or limbs

(ray-finned fishes) maneuverable fins supported by rays

Actinistia Ancient lineage of aquatic lobe-fins still surviving
Amniotes: amniotic egg, rib cage ventilation
Tetrapods: four limbs, neck, fused

(coelacanths) in Indian Ocean

Dipnoi Freshwater lobe-fins with both lungs and gills; sister
(lungfishes) group of tetrapods
Amphibia Have four limbs descended from modified fins; most
(salamanders, have moist skin that functions in gas exchange; many
frogs, caecilians) live both in water (as larvae) and on land (as adults)

Reptilia One of two groups of living amniotes; have amniotic

pelvic girdle

(tuataras, lizards eggs and rib cage ventilation, key adaptations for life
and snakes, turtles, on land
crocodilians, birds)

Mammalia Evolved from synapsid ancestors; include egg-laying

(monotremes, monotremes (echidnas, platypus); pouched marsupials
marsupials, (such as kangaroos, opossums); and eutherians
eutherians) (placental mammals, such as rodents, primates)
FUENTE:

Campbell Biology ninth Edition,

2011