Chapt 31

643
Animals: Part II
Chapter Concepts
31.1 Echinoderms
Echinoderms (e.g., starsh) and chordates (e.g.,
vertebrates) are both deuterostomes. 644
In deuterostomes, the second embryonic opening
becomes the mouth, and the coelom develops by
outpocketing from the primitive gut. 644
Echinoderms have radial symmetry and a
unique water vascular system for locomotion.
645
31.2 Chordates
All chordates have a notochord, a dorsal hollow
nerve cord, and pharyngeal pouches sometime
during their life history; in vertebrates, the
notochord is replaced by the vertebral column.
646
31.3 Vertebrates
There are three groups of shes. One group is
jawless, but the other two groupsthe
cartilaginous and bony shes (ray-nned and
lobe-nned)have jaws. 650
Amphibians (e.g., frogs and salamanders)
evolved from lobe-nned shes and have limbs,
an adaptation for locomotion on land. 651
The shelled egg of reptiles, which contains
extraembryonic membranes, is an adaptation for
reproduction on land. 653
Both birds, which can y, and mammals, which
have hair and mammary glands, evolved from
reptiles and are able to maintain a constant body
temperature. 655
31.4 Human Evolution
Primates (e.g., prosimians, monkeys, apes, and
humans) are mammals adapted to living in trees.
659
Human evolution diverged from ape evolution
in Africa about 4 million years ago. The
australopithecines were the rst hominids. 660
Homo habilis could make tools; Homo erectus
migrated out of Africa and was a big game
hunter. 661
Cro-Magnon is the name given to modern
humans who made sophisticated tools and
denitely had a culture. 664
Over half of the vertebrates alive today are shes. They come in
all sizes and shapesa coral reef is home for these lemon
buttery sh, Chaetodon miliaris.
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31.1 Echinoderms
In echinoderms, as well as
chordates, the second embry-
onic opening becomes the
mouth. Therefore, echino-
derms and chordates are
called deuterostomes. In these
animals, the coelom forms by
outpocketing of the primitive
gut, and therefore they are
also called enterocoelomates.
Echinoderms and close rela-
tives of chordates have a dipleurula larva (bands of cilia
placed as shown in Fig. 30.10).
Because the chordates include the vertebrates (e.g.,
human beings), it may seem surprising to learn that chor-
dates are most closely related to the echinoderms (e.g., sea
stars) that lack those features we associate with vertebrates.
For example, the echinoderms are often radially and not
bilaterally symmetrical. However, their larva is a free-
swimming lter feeder with bilateral symmetry. Metamor-
phosis results in the radially symmetrical adult.
Characteristics of Echinoderms
Echinoderms (phylum Echinodermata, about 6,000 species),
a diverse group of marine animals, have an endoskeleton (in-
ternal skeleton) consisting of spine-bearing, calcium-rich
plates. The spines, which stick out through their delicate
skin, account for their name. The echinoderms include only
marine animalssea stars, sea urchins, sea cucumbers,
feather stars, sea lilies, and sand dollars (Fig. 31.1).
644 Part 6 Evolution and Diversity 31-2
A
s Wendy waited in a camouaged blind for her next
target, the wildlife photojournalist thought about her
recent assignments. Last month, she had donned
scuba gear to photograph the remarkably ugly hagsh, and
also worked underwater in a cage to capture pictures of the
erce hammerhead shark. Her cameras had documented al-
ligators chasing down prey and turtles returning to the ocean
after burying their eggs on the seashore. Using a hang glider,
Wendy had even photographed a bald eagle ying around its
nest on the side of a steep mountain. Other jobs had her
track kangaroos in Australia and apes in Africa. Today, she
was waiting for a lion to attack the herd of antelopes at the
watering hole near her blind.
From sh to birds, reptiles to mammals, the vertebrate
world offers Wendy a seemingly endless number of photo-
graphic subjects. Many of those creatures will make an ap-
pearance in this chapter. We'll also detail how, through
evolution, one line of descent gave rise to the uniquely intel-
ligent species called Homo sapiens sapiens, otherwise
known as modern humans.
Figure 31.1 Echinoderm diversity.
a. Sea urchins have large, colored, external spines for protection.
b. Sea cucumbers look like a cucumber; they lack arms but have
tentaclelike tube feet with suckers around the mouth. c. A feather
star extends its arms to strain suspended food particles from the sea.
a. Purple sea urchin, Strongylocentrotus
b. Sea cucumber, Parastichopus
c. Feather star, Comanthus
Ancestral protists
coelomates
deutero-
stomes
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Sea Stars
Sea stars are commonly found along rocky coasts where
they feed on clams, oysters, and other bivalve mollusks. The
ve-rayed body has an oral, or mouth, side (the underside)
and an aboral, or anus, side (the upper side) (Fig. 31.2). Var-
ious structures project through the body wall: (1) spines
from the endoskeletal plates offer some protection; (2) pin-
cerlike structures called pedicellariae keep the surface free
of small particles; and (3) skin gills, tiny ngerlike exten-
sions of the skin, are used for gas exchange. On the oral sur-
face, each arm has a groove lined by small tube feet.
To feed, a sea star positions itself over a bivalve and at-
taches some of its tube feet to each side of the shell. By work-
ing its tube feet in alternation, it pulls the shell open. Avery
small crack is enough for the sea star to evert its cardiac
stomach and push it through the crack, so that it contacts the
soft parts of the bivalve. The stomach secretes enzymes, and
digestion begins even while the bivalve is attempting to
close its shell. Later, partly digested food is taken into the sea
stars body, where digestion continues in the pyloric stom-
ach using enzymes from the digestive glands found in each
arm. Ashort intestine opens at the anus on the aboral side.
In each arm, the well-developed coelomic cavity con-
tains not only a pair of digestive glands, but also gonads (ei-
ther male or female), which open on the aboral surface by
very small pores. The nervous system consists of a central
nerve ring that gives off radial nerves in each arm. A light-
sensitive eyespot is at the tip of each arm. Sea stars are capa-
ble of coordinated but slow responses and body movements.
Locomotion depends on the water vascular system.
Water enters this system through a structure on the aboral
side called the sieve plate, or madreporite. From there it
passes down a stone canal into a ring canal, which sur-
rounds the mouth. The ring canal gives off a radial canal in
each arm. From the radial canals, water enters the ampullae.
Contraction of the ampulla forces water into the tube foot,
expanding it. When the foot touches a surface, the center is
withdrawn, giving it suction so that it can adhere to the sur-
face. By alternating the expansion and contraction of the
tube feet, a sea star moves slowly along.
Echinoderms dont have a respiratory, excretory, or cir-
culatory system. Fluids within the coelomic cavity and the
water vascular system carry out many of these functions.
For example, gas exchange occurs across the skin gills and
the tube feet. Nitrogenous wastes diffuse through the
coelomic uid and the body wall. Cilia on the peritoneum
lining the coelom keep the coelomic uid moving.
Sea stars reproduce asexually and sexually. If the body is
fragmented, each fragment can regenerate a whole animal.
Fishermen who try to get rid of sea stars by cutting them up
and tossing them overboard are merely propagating more
sea stars! Sea stars spawn, releasing either eggs or sperm.
The dipleurula larva is bilateral and metamorphoses to be-
come the radially symmetrical adult.
Echinoderms have a well-developed coelom and
internal organs despite being radially symmetrical.
Spines project from their endoskeleton, and there
is a unique water vascular system.
Chapter 31 Animals: Part II 645 31-3
Figure 31.2 Sea star anatomy and behavior.
a. A sea star uses the suction of its tube feet to open a clam, its primary source of food. b. Each arm of a sea star contains digestive glands,
gonads, and portions of the water vascular system. This system (colored yellow) terminates in tube feet.
pedicellaria
radial canal
tube feet
ampulla
spine
gonads
digestive
gland
skin gill
central disk
anus
cardiac stomach pyloric stomach sieve plate
endoskeletal plates
arm
eyespot
tube feet
coelomic
cavity
a.
b.
31.2 Chordates
To be considered a chordate,
(phylum Chordata, about
45,000 species) an animal must
have the four basic character-
istics listed below at some time
during its life history:
1. Adorsal supporting rod
called a notochord. The
notochord is located just
below the nerve cord.
Vertebrates have an
embryonic notochord that is replaced by the vertebral
column during development.
2. Adorsal hollow nerve cord. By hollow, it is meant that the
cord contains a canal lled with uid. In vertebrates,
the nerve cord, more often called the spinal cord, is
protected by the vertebrae.
3. Pharyngeal pouches. These are seen only during
embryonic development in most vertebrates. In the
invertebrate chordates, the shes, and some amphibian
larvae, the pharyngeal pouches become functioning
gills. Water passing into the mouth and the pharynx
goes through the gill slits, which are supported by gill
arches. In terrestrial vertebrates which breathe by lungs,
the pouches are modied for various purposes. In
humans, the rst pair of pouches become the auditory
tubes. The second pair become the tonsils, while the
third and fourth pairs become the thymus gland and
the parathyroids.
4. Atail that extends beyond the anus and is therefore
called a post-anal tail.
notochord
post-
anal
tail
pharyngeal pouches
dorsal hollow
nerve cord
Ancestral protists
coelomates
deutero-
stomes
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The Deuterostomes* The second embryonic opening is associated with the mouth; enterocoelomates; dipleurula larva.
Phylum Echinodermata (echinoderms) Radial symmetry; endoskeleton of spine-bearing plates; water vascular system
with tube feet; about 6,000 species.
Phylum Chordata (chordates) Notochord, dorsal hollow nerve cord, pharyngeal pouches; about 45,000 species.
Subphylum Cephalochordata (chordates) Marine shlike animals with three chordate characteristics as adults.
Subphylum Urochordata (urochordates) Larva free-swimming with three chordate characteristics; adults sessile lter feeders with
plentiful gill slits; about 1,250 species.
Subphylum Vertebrata (vertebrates) Notochord replaced by vertebrae that protect the nerve cord; skull that protects the brain;
segmented with jointed appendages; about 43,700 species.
Superclass Agnatha (jawless shes) Marine and freshwater shes; lack jaws and paired appendages; notochord; about 63 species.
Superclass Gnathostomata Hinged jaws; paired appendages.
(jawed shes and tetrapods)
Class Chondrichthyes Marine cartilaginous shes; lack operculum and swim bladder; tail n usually
(cartilaginous shes) asymmetrical; about 850 species.
Class Osteichthyes Marine and freshwater bony shes; operculum; swim bladder or lungs; tail n; usually
(bony shes) symmetrical; about 20,000 species.
Class Amphibia (amphibians) Tetrapod with nonamniote egg; nonscaly skin; three-chambered heart; ectothermic;
metamorphosis; about 3,900 species.
Class Reptilia (reptiles) Tetrapod with amniote egg; scaly skin; ectothermic; teeth not differentiated typically; about
6,000 species.
Class Aves (birds) Tetrapod with feathers; bipedal with wings; double circulation; endothermic; about 9,000 species.
Class Mammalia (mammals) Tetrapods with hair, mammary glands; double circulation; endothermic; teeth
differentiated. Monotremes, marsupials, placental mammals; about 4,500 species.
Order Primates Large cerebral hemispheres; mostly tree dwellers; opposable thumb and at nails; about 233
species
*Not in the classication of organisms, but added here for clarity.
Figure 31.3 Evolutionary tree of chordates.
Each of the highlighted features is an evolved characteristic that is shared by the classes beyond this point.
jaws
lungs
limbs
amnion
mammary
glands, hair
feathers
vertebrae
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Ancestral chordates
Invertebrate
Chordates
There are a few invertebrate
chordates in which the noto-
chord is never replaced by the
vertebral column.
Lancelets, formerly re-
ferred to as amphioxus, (sub-
phylum Cephalochordata,
about 23 species), are now
classied in the genus
Branchiostoma. These marine
chordates, which are only a few centimeters long, are named
for their resemblance to a lanceta small, two-edged surgi-
cal knife (Fig. 31.4). Lancelets are found in the shallow water
along most coasts where they usually lie partly buried in
sandy or muddy substrates with only their anterior mouth
and gill apparatus exposed. They feed on microscopic parti-
cles ltered out of the constant stream of water that enters
the mouth and exits through the gill slits.
Lancelets retain the four chordate characteristics as an adult.
In addition, segmentation is present, as witnessed by the fact
that the muscles are segmentally arranged and the dorsal
hollow nerve cord has periodic branches.
Tunicates, or sea squirts (subphylum Urochordata,
about 1,250 species), live on the ocean oor and take their
name from a tunic that makes the adults look like thick-
walled, squat sacs. They are also called sea squirts because
they squirt out water from one of their siphons when dis-
turbed. The tunicate larva is bilaterally symmetrical and has
the three chordate characteristics. Metamorphosis produces
the sessile adult with an incurrent and excurrent siphon
(Fig. 31.5).
The pharynx is lined by numerous cilia whose beating
creates a current of water that moves into the pharynx and
out the numerous gill slits, the only chordate characteristic
that remains in the adult. Microscopic particles adhere to a
mucous secretion and are eaten.
Is it possible that the tunicates are directly related to the
vertebrates? It has been suggested that a larva with the four
chordate characteristics may have become sexually mature
without developing the other adult tunicate characteristics.
Then it may have evolved into a shlike vertebrate.
The invertebrate chordates include the tunicates
and the lanceletsa lancelet is the best example
of a chordate that possesses the three chordate
characteristics as an adult.
tail
dorsal fin
anus
ventral fin
atriopore
atrium
gill slits
gill bar
dorsal hollow
nerve cord
notochord
oral hood with tentacles
pharynx
Figure 31.5 Anatomy of a tunicate, Halocynthia.
Note that the only chordate characteristic remaining in the adult is
gill slits.
Figure 31.4 Habitat and anatomy of a lancelet,
Branchiostoma.
Water enters the mouth and exits at the atripore after passing
through the gill slits. Lancelets are lter feeders.
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Ancestral chordates
31.3 VertebratesM
At some time in their life
history, vertebrates (sub-
phylum Vertebrata, about
43,700 species) have all three
chordate characteristics. The
embryonic notochord, how-
ever, is generally replaced
by a vertebral column com-
posed of individual verte-
brae. The vertebral column,
which is a part of the exible
but strong jointed endoskeleton, gives evidence that verte-
brates are segmented. The skeleton protects internal or-
gans and serves as a place of attachment for muscles.
Together the skeleton and muscles form a system that per-
mits rapid and efcient movement. Two pairs of appendages
are characteristic. The pectoral and pelvic fins of fish
evolved into the jointed appendages that allowed verte-
brates to move onto land.
The main axis of the endoskeleton consists of not only
the vertebral column, but also a skull that encloses and pro-
tects the brain. The high degree of cephalization is accompa-
nied by complex sense organs. The eyes develop as
outgrowths of the brain. The ears are primarily equilibrium
devices in aquatic vertebrates, but they also function as
sound-wave receivers in land vertebrates.
The evolution of jaws in vertebrates has allowed some
to take up the predatory way of life. Vertebrates have a com-
plete digestive tract and a large coelom. The circulatory sys-
tem is closed (the blood is contained entirely within blood
vessels). Vertebrates have an efcient means of obtaining
oxygen from water or air, as appropriate. The kidneys are
important excretory and water-regulating organs that con-
serve or rid the body of water as necessary. The sexes are
generally separate, and reproduction is usually sexual. The
evolution of the amnion allowed reproduction to take place
on land. Reptiles, birds and some mammals lay a shelled
egg. In placental mammals the development takes place
within the uterus of the female.
Figure 31.6 shows major milestones in the evolutionary
history of vertebrates: the evolution of jaws, limbs, and the
amnion, an extraembryonic membrane which is rst seen in
the shelled amniote egg.
Vertebrates are distinguished in particular by:
living endoskeleton
closed circulatory system
paired appendages
efcient respiration and excretion
high degree of cephalization
In short, vertebrates are adapted to an active
lifestyle.
Figure 31.6 Milestones in vertebrate evolution.
a. The evolution of jaws in shes allows animals to be predators and
feed off other animals. b. The evolution of limbs in amphibians is
adaptive for locomotion on land. c. The evolution of a shelled egg in
reptiles is adaptive for reproduction on land.
a. Great white shark, Carcharodon
vertebrae
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Ancestral chordates
b. Spotted salamander, Ambystoma
c. Veiled chameleons, Chamaeleo
Fishes
The rst vertebrates were
shlike, and today there are
three living classes: jawless
shes, cartilaginous shes,
and the bony shes.
Jawless Fishes
Living representatives of the
jawless shes (superclass
Agnatha, about 63 species)
are cylindrical and up to a
meter long. They have
smooth, scaleless skin and no jaws or paired ns. There are
two groups of living jawless shes: hagshes and lampreys.
The hagshes are scavengers, feeding mainly on dead shes,
while some lampreys are parasitic. When parasitic, the
round mouth of the lamprey serves as a sucker. The lamprey
attaches itself to another sh and taps into its circulatory
system. Water cannot move in through the mouth and out
over the gills as is common in all other shes. Instead, water
moves in and out through the gill openings.
Fishes with Jaws
All of the other shes have jaws, tooth-bearing bones of the
head. Jaws are believed to have evolved from the rst pair of
gill arches. The second pair of gill arches became support
structures for jaws. The presence of jaws is an adaptation to
a predatory way of life.
Cartilaginous Fishes Cartilaginous shes (class Chon-
drichthyes, about 850 species) are the sharks, the rays, and
the skates, which have skeletons of cartilage instead of bone.
The small dogsh shark is often dissected in biology labora-
tories. One of the most dangerous sharks inhabiting both
tropical and temperate waters is the hammerhead shark.
The largest sharks, the whale sharks, feed on small shes
and marine invertebrates and do not attack humans. Skates
and rays are rather at shes that live partly buried in the
sand and feed on mussels and clams.
Three well-developed senses enable sharks and rays
to detect their prey. They have the ability to sense electric
currents in watereven those generated by the muscle
movements of animals. They have a lateral line system, a
series of pressure-sensitive cells that lie within canals
along both sides of the body, which can sense pressure
caused by a sh or other animal swimming nearby. They
also have a keen sense of smell; the part of the brain asso-
ciated with this sense is twice as large as the other parts.
Sharks can detect about one drop of blood in 115 liters (25
gallons) of water.
Bony Fishes Bony shes (class Osteichthyes, about 20,000
species) are by far the most numerous and diverse of all the
vertebrates. Most of the shes we eat, such as perch, trout,
salmon, and haddock are a type of bony sh called ray-nned
shes. Ray-nned shes are the most successful and diverse
of all the vertebrates. Some, like herrings, are lter feeders;
others, like trout, are opportunists, and still others are
predaceous carnivores, like piranhas and barracudas.
caudal fin anal fin second dorsal fin first dorsal fin
lateral line
intestine
kidney
swim bladder
stomach
muscle
bony vertebra
brain
nostril
gills
heart
liver
gallbladder
pelvic fin pectoral fin
scales
operculum
gonad
Figure 31.7 External and internal anatomy of a ray-nned sh called a soldiersh, Myripristis.
jaws
lungs
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Ancestral chordates
Bony shes have a swim bladder, which usually serves as
a buoyancy organ (Fig. 31.7). By secreting gases into the
bladder or by absorbing gases from it, these shes can
change their density and thus go up or down in the water.
Ray-nned refers to the fact that the paired ns, which are
paddlelike processes used in balancing and propelling the
body, are thin and supported by bony rays.
Fishes are adapted to life in the water. Usually sperm
and eggs are shed into the water, where fertilization occurs.
The zygote develops into a swimming larva, which can fend
for itself until it develops into the adult form. The stream-
lined shape, ns, and muscle action of most bony shes are
all suited to locomotion in the water. Their skin is covered by
bony scales which protect the body but do not prevent water
loss. When shes respire, the gills are kept continuously
moist by the passage of water through the mouth and out
the gill slits. As the water passes over the gills, oxygen is ab-
sorbed by blood and carbon dioxide is given off. Fishes have
a single-circuit circulatory system. The heart is a simple
pump, and the blood ows through the chambers, including
a nondivided atrium and ventricle, to the gills. Oxygenated
blood leaves the gills and goes to the body proper, eventu-
ally returning to the heart for recirculation.
Another type of bony sh called the lobe-nned shes
evolved into the amphibians. These shes not only had
eshy appendages that could be adapted to land locomo-
tion; most also had a lung, which was used for respiration. A
type of lobe-nned sh called the coelacanth, which exists
today, is the only living fossil among the shes. The coela-
canth, however, does not have a lung.
Most shes today are ray-nned shes. They
have the following characteristics:
bony skeleton and scales
swim bladder
two-chambered heart
paired ns
jaws
gills
Amphibians
Amphibians (class Amphibia,
about 3,900 species), whose
class name means living on
both land and in the water, are
represented today by frogs,
toads, newts, and salaman-
ders. Aside from being
tetrapods (presence of two sets
of paired limbs) and appropri-
ate modications of the gir-
dles, amphibians have other
features not seen in bony
shes. Their eyes have eyelids for keeping eyes moist, ears
adapted to picking up sound waves, and a voice-producing
larynx. The brain is larger than that of a sh. Adult amphibians
usually have small lungs. Air enters the mouth by way of nos-
trils, and when the oor of the mouth is raised, air is forced
into the relatively small lungs. Respiration is supplemented by
gas exchange through the smooth, moist, and glandular skin.
The amphibian heart has a divided atrium but a single ventri-
cle. The right atrium receives blood that has little oxygen from
the body proper, and the left atrium receives blood that has
much oxygen from the lungs. These two types of blood are
partially mixed in the single ventricle (see page 654). Mixed
blood is then sent to all parts of the body; some is sent to the
skin, where it is further oxygenated.
Most members of this group lead an amphibious lifethat
is, the larval stage lives in the water and the adult stage lives on
the land. The adult usually returns to the water, however, to re-
produce. Figure 31.8 illustrates how the frog tadpole undergoes
metamorphosis into the adult before taking up life on the land.
These features in particular distinguish
amphibians:
usually tetrapods usually lungs in adults
mostly metamorphosis smooth, moist skin
three-chambered heart
Figure 31.8 Frog metamorphosis.
During this time, the animal changes from an aquatic to a terrestrial organism.
a. Tadpoles hatch b. Tadpole respires with
external gills
c. Front and hind legs are
present
d. Frog respires with
lungs
limbs
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Ancestral chordates
Todays paleontologists are setting the record straight about di-
nosaurs. Because dinosaurs are classied as reptiles, it is as-
sumed that they must have had the characteristics of todays
reptiles. They must have been ectothermic, slow moving, and
antisocial, right? Wrong!
First of all, not all dinosaurs were great lumbering beasts.
Many dinosaurs were less than 1 meter (3 feet) long and their
tracks indicate they moved at a steady pace. These dinosaurs
stood on two legs that were positioned directly under the body.
Perhaps they were as agile as ostriches, which are famous for
their great speed.
Dinosaurs may have been endothermic. Could they have
competed successfully with the preevolving mam-
mals otherwise? They must have been able to hunt
prey and escape from predators as well as mammals,
which are known to be active because of their high
rate of metabolism. Some argue that ectothermic ani-
mals have little endurance and cannot keep up. They
also believe that the bone structure of dinosaurs indi-
cates they were endothermic.
Dinosaurs cared for their young much like birds
do today. In Montana, paleontologist Jack Horner
has studied fossilized nests complete with eggs, em-
bryos, and nestlings (Fig. 31A). The nests are about
7.5 meters (24.6 feet) apart, the space needed for the
length of an adult parent. About 20 eggs are laid in
neatly arranged circles and may have been covered
with decaying vegetation to keep them warm. Many
contain the bones of juveniles as much as a meter
long. It would seem then that baby dinosaurs re-
mained in the nest to be fed by their parents. They
must have obtained this size within a relatively short
period of time, again indicating that dinosaurs were
endothermic. Ectothermic animals grow slowly and
take a long time to reach this size.
Dinosaurs were also social! An enormous herd of
dinosaurs found by Horner and colleagues is esti-
mated to have nearly 30 million bones, representing
10,000 animals in one area measuring about 1.6
square miles. Most likely, the herd kept on the move
in order to be assured of an adequate food supply,
which consisted of owering plants that could be
stripped one season and grow back the next season.
The fossilized herd is covered by volcanic ash, sug-
gesting that the dinosaurs died following a volcanic
eruption.
Some dinosaurs, such as the duck-billed di-
nosaurs and horned dinosaurs, have a skull crest.
How might it have functioned? Perhaps it was a res-
onating chamber, used when dinosaurs communi-
cated with one another. Or, as with modern horned animals that
live in large groups, the males could have used the skull crest in
combat to establish dominance.
Some zoologists do not agree with the assertions of paleon-
tologists. They believe that dinosaurs did have the characteris-
tics of todays reptiles, and this doesnt make them in any way
inferior to endothermic mammals. Ectothermic animals can
wait out cold weather conditions, while endothermic animals
must always venture forth to nd food, no matter what the
conditions.
652
Real Dinosaurs, Stand Up!
Figure 31A Behavior of dinosaurs.
a. Nest of fossil dinosaur eggs found in Montana, dating from the Cretaceous
period. b. Bones of a hatchling (about 50 cm [20 inches]) found in the nest.
These dinosaurs have been named Maiosaura, which means good mother
lizard in Greek.
b.
a.
Reptiles
Reptiles (class Reptilia, about
6,000 species) diversied and
were abundant some 245 to
66 millions of years ago.
Aside from those that lived
on to become the living rep-
tiles of today, there were the
mammal-like reptiles and the
dinosaurs, which became ex-
tinct. Some dinosaurs are re-
membered for their great size.
Brachiosaurus, a herbivore, was
about 23 meters (75 feet) long and about 17 meters (56 feet)
tall. Tyrannosaurus rex, a carnivore, was 5 meters (16 feet) tall
when standing on its hind legs. A bipedal stance was
preadaptive for the evolution of wings in birds. Some say
birds are actually living dinosaurs; new studies on the be-
havior of dinosaurs discussed in the accompanying reading
give credence to this hypothesis.
The reptiles living today are mainly turtles, alligators,
snakes, and lizards. The body is covered with hard, kera-
tinized scales, which protect the animal from desiccation and
from predators. Another adaptation for a land existence is
the manner in which snakes use their tongue as a sense or-
gan (Fig. 31.9). Reptiles have well-developed lungs enclosed
by a protective rib cage. When the rib cage expands, the
lungs expand and air rushes in. The creation of a partial vac-
uum establishes a negative pressure, which causes air to
rush into the lungs. The atrium of the heart is always sepa-
rated into right and left chambers, but division of the ventri-
cle varies. An interventricular septum is incomplete in
certain species; therefore, there is some exchange of oxy-
genated and deoxygenated blood between the ventricles.
Perhaps the most outstanding adaptation of the reptiles
is that they have a means of reproduction suitable to a land
existence. The penis of the male passes sperm directly to the
female. Fertilization is internal, and the female lays leathery,
exible, shelled eggs. The amniote egg made development on
land possible and eliminated the need for a swimming-
larval stage during development. It provides the developing
embryo with atmospheric oxygen, food, and water; it re-
moves nitrogenous wastes; and it protects the embryo from
drying out and from mechanical injury. This is accom-
plished by the presence of extraembryonic membranes
(Fig. 31.10).
Fishes, amphibians, and reptiles are ectothermic. Their
body temperature matches the temperature of the external
environment. If it is cold externally, they are cold internally;
if it is hot externally, they are hot internally. Reptiles try to
regulate body temperatures by exposing themselves to the
sun if they need warmth or by hiding in the shadows if they
need cooling off. This works reasonably well in most areas
of the world.
These features in particular distinguish reptiles:
usually tetrapods shelled egg
lungs with expandable dry, scaly skin
rib cage
a.
albumin
air
space
yolk sac
chorion
amnion
embryo
egg shell
allantois
Figure 31.9 The tongue as a sense organ.
Snakes wave a forked tongue in the air to collect chemical molecules
which are brought back into the mouth and then delivered to an
organ in the roof of the mouth. Analyzed chemicals help the snake
trail a prey animal, recognize a predator, or nd a mate.
Figure 31.10 The reptilian egg allows reproduction on land.
a. Baby American crocodile, Crocodylus, hatching out of its shell.
Note that the shell is leathery and exible, not brittle like birds eggs.
b. Inside the egg, the embryo is surrounded by membranes. The
chorion aids gas exchange, the yolk sac provides nutrients, the
allantois stores waste, and the amnion encloses a uid that prevents
drying out and provides protection.
amnion
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Ancestral chordates
b.
Figure 31.11 Anatomy of a bird.
a. Birds have a large, keeled sternum to which ight muscles attach. Bird bones are strong but weigh very little because they contain air cavities.
In feathers, a hollow central shaft gives off barbs and barbules, which interlock in a latticelike array. b. In birds and mammals, the heart has four
chambers and sends only blood that has little oxygen to the lungs and blood that has much oxygen to the body. In amphibians, the heart has two
chambers and there is some mixture of oxygenated and deoxygenated blood. In shes, a single-looped system utilizes a two-chambered heart.
mandible
skull
radius
ulna
humerus
pectoral
girdle
sternum
with keel
ribs
pelvic
girdle
femur
barb
barbule
shaft
a.
b.
gill
capillaries
gill
circulation
pulmonary
capillaries
pulmonary
circulation
pulmonary
circulation
pulmonary
capillaries
Fish Amphibian
Key:
oxygenated blood
deoxygenated blood
mixed blood
Bird and mammal
systemic
circulation
systemic
capillaries
systemic
capillaries
systemic
capillaries
systemic
circulation
systemic
circulation
ventricle
atrium
aorta
aorta
aorta
right
atrium
left
ventricle
Birds have a four-
chambered heart
and separate
pulmonary and
systemic systems.
A feather is a single unit. The
barbs that project from the
shaft have barbules that
hook together.
Feathers not only
allow flight, they
also help maintain a
relatively high body
temperature.
VISUAL FOCUS
ature is constant because they generate and maintain meta-
bolic heat. This may be associated with their efcient ner-
vous, respiratory, and circulatory systems. Also, their
feathers provide insulation. Birds have no bladder and ex-
crete uric acid in a semidry state.
Birds have well-developed brains; the enlarged portion
seems to be the area responsible for instinctive behavior. A
ritualized courtship often precedes mating. Birds practice
internal fertilization and lay hard-shelled eggs. Many newly
hatched birds require parental care before they are able to y
away and seek food for themselves. Aremarkable aspect of
bird behavior is the seasonal migration of many species over
very long distances. Birds navigate by day and night, and
whether its sunny or cloudy, by using the sun and stars and
even the earths magnetic eld to guide them.
There are many orders of birds, including birds that are
ightless (ostriches), web-footed (penguins), divers (loons),
sh eaters (pelicans), waders (amingos), broad billed
(ducks), birds of prey (hawks), vegetarians (fowl, e.g., chick-
ens and turkeys), shorebirds (sandpipers), nocturnal (owls),
small (hummingbirds), large (condors), and songbirds, the
most familiar of the birds.
These features in particular distinguish birds:
usually wings for ying feathers
hard-shelled egg air sacs
four-chambered heart endothermic
Birds
Birds (class Aves, about 9,000
species) are characterized by
the presence of feathers (Fig.
31.11), which are modied
reptilian scales. (Perhaps you
have noticed that there are
scales on the legs of a chicken.)
Nearly every anatomical
feature of a bird can be re-
lated to its ability to y. The
anterior pair of appendages
(wings) are adapted for ight;
the posterior are variously modied, depending on the type
of bird. Some are adapted for swimming, some for running,
and some for perching on limbs. The breastbone is well de-
veloped and has a ridge, the keel, to which the ight muscles
are attached (Fig. 31.11). Respiration is efcient since the lob-
ular lungs form anterior and posterior air sacs. The presence of
these sacs means that the air circulates one way through the
lungs and there is a continuous exchange of gases across re-
spiratory tissues (Fig. 31.12). Another benet of air sacs is
that they lighten the body and aid ying.
Birds have a four-chambered heart which completely
separates blood with much oxygen from blood that has little
oxygen (see page 654). Blood that has a relatively high con-
centration of oxygen is sent under pressure to the muscles.
Birds are endothermic. Like mammals, their internal temper-
trachea
anterior
air sacs
a.
lung
posterior
air sacs
Inspiration
lung
anterior
air sacs posterior
air sacs
trachea
Expiration
Expiration Expiration
b.
Figure 31.12 How birds breathe.
a. Birds have a system of air sacs in their bones. b. When a bird inhales, air enters the posterior air sacs, and when a bird exhales, air moves
through the lungs to the anterior air sacs before exiting the trachea. This one-way ow of air is efcient and allows more oxygen to be removed
from one breath of air.
feathers
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Ancestral chordates
Mammals
Mammals also evolved
from the reptiles. These first
mammals were small, about
the size of mice. All the time
the dinosaurs flourished
(165 million years ago),
mammals were a minor
group that changed little.
Some of the earliest mam-
malian groups, represented
today by the monotremes
and marsupials, are not
abundant today. The placental mammals that evolved
later went on to live in many habitats, including air, land,
and sea habitats.
The chief characteristics of mammals (class Mammalia,
about 4,500 species) are the possession of hair and milk-
producing mammary glands. They are almost all endother-
mic and maintain a constant internal temperature. Many of
the adaptations of mammals are related to temperature con-
trol. Hair, for example, provides insulation against heat loss
and allows mammals to be active, even in cold weather. Like
birds, mammals have efcient respiratory and circulatory
systems, which assure a ready oxygen supply to muscles
whose contraction produces body heat. Like birds, mam-
mals have a double-loop circulation and a four-chambered
heart (see Fig. 31.11b).
Mammary glands enable females to feed (nurse) their
young without leaving them to nd food. Nursing also cre-
ates a bond between mother and offspring that helps en-
sure parental care while the young are helpless. In most
mammals, the young are born alive after a period of devel-
opment in the uterus, a part of the female reproductive
tract. Internal development shelters the young and allows
the female to move actively about while the young are ma-
turing. Mammals are classified according to how they
reproduce.
Monotremes
Monotremes are mammals that, like birds, have a cloaca, a
terminal region of the digestive tract serving as a common
chamber for digestive, excretory wastes, and sex cells. They
also lay hard-shelled amniote eggs. They are represented by
the duckbill platypus and the spiny anteater, both of which
are found in Australia (Fig. 31.13a). The female duckbill
platypus lays her eggs in a burrow in the ground. She incu-
bates the eggs and, after hatching, the young lick up milk
that seeps from modied sweat glands on the abdomen of
a. Duckbill platypus, Ornithorhynchus
b. Koala, Phascolarctos
Figure 31.13 Three types of mammals.
a. The duckbill platypus is a monotreme, which lays shelled eggs.
b. The koala is a marsupial, whose young are born immature and
complete their development within the mothers pouch. c. The white-
tailed deer is a placental mammal, whose young develop within a
uterus.
hair,
mammary
glands
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Ancestral chordates
both males and females. The spiny anteater has a pouch on
the belly side formed by swollen mammary glands and lon-
gitudinal muscle. The egg moves from the cloaca to this
pouch, where hatching takes place, and the young remain
for about 53 days. Then they stay in a burrow, where the
mother periodically visits and nurses them.
Marsupials
The young of marsupials begin their development inside
the females body, but they are born in a very immature con-
dition. Newborns crawl up into a pouch on their mothers
abdomen. Inside the pouch, they attach to nipples of mam-
mary glands and continue to develop. Frequently, more are
born than can be accommodated by the number of nipples,
and its rst come, rst served.
Today, marsupial mammals are found mainly in Aus-
tralia, but also in Central and South America. They could not
compete against placental mammals, and only a few marsu-
pials, such as the American opossum, are found in North
America. In Australia, marsupials underwent adaptive radi-
ation for several million years without competition from
placental mammals, which arrived there only recently.
Among the herbivorous marsupials, koalas are tree-
climbing browsers (Fig. 31.13b) and kangaroos are grazers.
The Tasmanian wolf or tiger, thought to be extinct, was a
carnivorous marsupial about the size of a collie dog.
Placental Mammals
The vast majority of living mammals are placental mam-
mals (Fig. 31.13c). In these mammals, the extraembryonic
membranes found in reptiles (see Fig. 31.11) have been mod-
ied for internal development within the uterus of the fe-
male. The chorion contributes to the fetal portion of the
placenta, while a part of the uterine wall contributes to the
maternal portion. Here, nutrients, oxygen, and waste are ex-
changed between fetal and maternal blood.
Mammals are adapted to life on land and have limbs
that allow them to move rapidly. In fact, an evaluation of
mammalian features leads us to the obvious conclusion that
they lead active lives. The brain is well developed; the lungs
are expanded not only by the action of the rib cage but also
by the contraction of the diaphragm, a horizontal muscle that
divides the thoracic cavity from the abdominal cavity; and
the heart has four chambers. The internal temperature is con-
stant, and hair, when abundant, helps to insulate the body.
The mammalian brain is enlarged due to the expansion of
the foremost partthe cerebral hemispheres (see Figure
17.12). These have become convoluted and have expanded to
such a degree that they hide many other parts of the brain
from view. The brain is not fully developed for some time af-
ter birth, and there is a long period of dependency on the par-
ents, during which the young learn to take care of themselves.
Mammals have differentiated teeth. Typically, in the
front, the incisors and canine teeth have cutting edges for
capturing and killing prey. On the sides, the premolars and
molars chew food. Classication of placental mammals is
based on methods of obtaining food and mode of locomo-
tion. For example, bats (order Chiroptera) have membra-
nous wings supported by digits; horses (order
Perissodactyla) have long, hoofed legs; and whales (order
Cetacea) have paddlelike forelimbs. The specic shape and
size of the teeth may be associated with whether the mam-
mal is an herbivore (eats vegetation), a carnivore (eats meat),
or an omnivore (eats both meat and vegetation). For exam-
ple, mice (order Rodentia) have continuously growing in-
cisors; horses (order Perissodactyla) have large, grinding
molars; and dogs (order Carnivora) have long canine teeth.
These features in particular distinguish placental
mammals:
body hair
differentiated teeth
infant dependency
constant internal temperature
mammary glands
well-developed brain
internal development
c. White-tailed deer, Odocoileus
Apes diverge from the
human line of descent.
Monkeys diverge from the
human line of descent.
Prosimians diverge from
the human line of descent.
Angiosperms evolve
and forests spread.
Mammalian ancestor
enters trees.
tarsier
capuchin
baboon
gibbon
gorilla
human
being
Epoch Prosimians
Holocene
Pleistocene
Pliocene
.01
2
6
Miocene
Oligocene
24
37
Eocene
Paleocene
58
66
C
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Y
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A
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(
M
Y
A
)
Anthropoids
Hominoids
New World Monkeys Old World Monkeys Asian Apes African Apes Hominids
Figure 31.14 Primate evolution.
There are at least four lines of evolution among the primates, which arose from mammalian insectivores: prosimians; monkeys, including New
World and Old World monkeys; apes, including Asian apes and African apes (orangutans, gorillas, and chimpanzees); and humans.
All of the animals depicted in Figure 31.14 are primates,
which are classied as shown in Table 31.1.
Primates
Primates (order Primates, about 180 species) belong to a
mammalian order in which the animals are adapted to liv-
ing in trees. The limbs are mobile, as are the hands, because
the thumb (and in nonhuman primates, the big toe as well)
is opposable; that is, the thumb can touch each of the other
ngers. Therefore, a primate can easily reach out and bring
food such as fruit to the mouth. When locomoting, tree
limbs can be grasped and released freely because nails have
replaced claws.
The sense of smell is of primary importance in animals
with a snout. In primates, the snout is shortened consider-
ably, allowing the eyes to move to the front of the head. The
stereoscopic vision (or depth perception) that results per-
mits primates to make accurate judgments about the dis-
tance and position of adjoining tree limbs.
Gestation is lengthy, allowing time for good forebrain
development; the visual portion of the brain is proportion-
ately large, as are those centers responsible for hearing and
touch. One birth at a time is the norm in primates; it is dif-
cult to care for several offspring while moving from limb to
limb. The juvenile period of dependency is extended, and
there is an emphasis on learned behavior and complex social
interactions.
The primate order contains two suborders: Prosimii and
Anthropoidea. The rst primates were the prosimians, a
term meaning premonkey. The prosimians are represented
today by several types of animals. Lemurs have a squirrel-
like appearance (Fig. 31.14), and tarsiers are curious mouse-
sized creatures with enormous eyes suitable for their
nocturnal way of life. The prosimians are believed to have
evolved from an insectivore-type mammal.
Monkeys, apes, and humans are all anthropoids (subor-
der Anthropoidea). There are two types of monkeys: New
World monkeys, which have long prehensile (grasping) tails
and at noses, and Old World monkeys, which lack such
tails and have protruding noses. Two of the well-known
New World monkeys are the spider monkey and the ca-
puchin, the organ grinders monkey. Some of the better
known Old World monkeys are now ground dwellers, such
as the baboon and the rhesus monkey.
Humans are more closely related to apes than to mon-
keys; only apes and humans are hominoids (superfamily
Hominoidea). There are four types of apes: gibbons, orangu-
tans, gorillas, and chimpanzees. Gibbons, the smallest of the
apes, have extremely long arms, which are specialized for
swinging between tree limbs. The orangutan is a large ape
but nevertheless spends a great deal of time in trees. In con-
trast, the gorilla, the largest of the apes, spends most of its
time on the ground. Chimpanzees, which are at home both
in the trees and on the ground, are the most humanlike of
the apes in appearance. Molecular data tell us that humans
are genetically very similar to chimpanzees and gorillas.
Nucleotide sequences in genes, amino acid sequences in
proteins, and immunological properties of various mole-
cules all indicate that we are even more closely related to
these apes than orangutans are related to them.
Humans can be distinguished from apes, however, by
locomotion and posture, dental features, and other charac-
teristics. When moving in a tree, a monkey is quadrupedal
(walks on all four limbs) and leaps from branch to branch.
In keeping with this method of locomotion, the vertebral
column is arched and the shoulder joint has limited mobil-
ity. In contrast to monkeys, apes swing from branch to
branch. Consistent with this type of locomotion, the verte-
bral column is straight and the shoulder joint is mobile.
Also, the arms are elongated. Humans are bipedalthat is,
they walk on two feet. Humans have an S-shaped vertebral
column, which provides a way to transmit the upper
bodys weight to the pelvis. The pelvis is stronger than that
of an ape because it is shorter, and the sacrum ts like a
keystone between the two pelvic bones. The broad pelvic
bones serve as attachments for muscles that maintain sta-
bility as rst one leg and then the other leaves the ground
while walking.
These characteristics especially distinguish
primates from other mammals:
opposable thumb (and in some cases, big toe)
extended period of parental care
emphasis on learned behavior
expanded forebrain
nails (not claws)
single birth
Phylum Chordata Invertebrate and vertebrate chordates
Subphylum Vertebrata Fish, amphibians, reptiles, birds,
mammals
Class Mammalia Monotremes, marsupials, placental
mammals
Order Primates Prosimians and anthropoids
Suborder Anthropoidea Monkeys, apes, and hominids
Superfamily Hominoidea Apes and hominids
Family Hominidae Australopithecines, Homo habilis,
Homo erectus, Homo sapiens
Table 31.1 Primate Classication
More than twenty years ago, a team led by Donald Jo-
hanson unearthed nearly 250 fossils of a hominid called Aus-
tralopithecus afarensis. A now-famous female skeleton dated
at about 3.4 MYA is known worldwide by its eld name,
Lucy. (The name derives from the Beatles song Lucy in the
Sky with Diamonds which they played in camp.) Although
her brain was about the size of a chimp (400 cc), the shapes
and relative proportions of her limbs indicate that Lucy
walked bipedally (Fig. 31.15). There are even footprints that
show how members of her species walked.
The australopithecines were sexually dimorphic. Lucy
was about four feet tall and weighed about 30 kilograms. In
contrast, the males of the species were ve feet tall and
weighed up to 45 kilograms. Some have speculated that
such size differences may indicate two separate species, but
in 1994 new nds, including a more complete skull (dubbed
the son of Lucy and dated at just about 3 MYA), conrmed
the opinion that the fossils belong to one species. Taking
into account their smaller body size, the relative brain size
is about one-third of ours and the jaw is heavy. The cheek
teeth are enormous, and in males, large canine teeth project
forward.
A. afarensis had descendants. After a period of stasis
that may have lasted a million years (from 3 to 2 MYA),
branching speciation occurred (Fig. 31.16). Therefore, in-
stead of thinking about descent in terms of a straight line, it
is far better to envision a bush. Some think there may have
been as many as ten species of hominids about 2 MYA in
Africa, but we will discuss only four of them, three of which
are australopithecines. A. africanus, which was rst named
in southern Africa by Raymond Dart in the 1920s, is a
gracile (slender) type. A. boisei is a robust form from eastern
Africa, and A. robustus is a similar form from southern
Africa. The robust forms have stronger jaws, larger attach-
ments for larger chewing muscles, and bigger grinding
teeth because they most likely fed on tougher foods than
the gracile form. The robust forms lived in drier habitats
where soft fruits and leaves would be harder to come by. In
both forms, the pelvis resembles that of Lucy but the hands
were more humanlike; possibly they were capable of mak-
ing tools. Both forms, which are believed to have eaten meat
at least occasionally, may have used the tools to process an-
imal carcasses.
At one time it was believed that the gracile australo-
pithecine form gave rise to the robust forms. However, in
1985, Alan Walker discovered a robust skull, called the black
skull, which was old enough (2.5 MYA) to be ancestral to the
robust forms we have been discussing. It has been given the
name A. aethiopicus.
Several species of australopithecines have been
identied. After a period of stasis, during which
only A. afarensis existed, branching speciation
occurred.
Hominids
The hominid line of descent begins with the australo-
pithecines, which evolved and diversied in eastern Africa.
It wasnt until the 1960s that paleontologists, following the
lead of the renowned couple Louis and Mary Leakey, began
to concentrate their efforts in eastern as opposed to southern
Africa. It has proven fruitful for paleontologists. One 1994
nd consisting of skull fragments, teeth, arm bones, and a
part of a childs lower jaw has been dated at 4.4 MYA (mil-
lions of years ago). Called A. ramidus, it is believed to repre-
sent an early stage in human evolution. In comparison to
later australopithecines, the canine teeth are larger, and the
skull is more like that of a chimpanzee. Also, this fossil did
not walk erect. However, a 1995 discovery, Australopithecus
anamensis, dated at just about 4 MYA, has jaws like those of
an ape but legs like those of humans.
Figure 31.15 Australopithecus afarensis.
Reconstruction of Lucy, who is believed to have walked erect.
A. afarensis was a dimorphic species, in which males were much
larger than the females. Lucy was four feet tall and weighed about 30
kilograms. A male was ve feet tall and weighed about 45 kilograms.
Homo habilis
The oldest fossils to be classied in the genus Homo are
known as Homo habilis (meaning handy man). His re-
mains, dated as early as 2 MYA, are often accompanied by
stone tools. Why is this hominid classied within our own
genus? H. habilis was smallabout the size of Lucybut
the brain, at 700 cc, is about 45% larger. In addition, certain
portions of the brain thought to be associated with speech
areas are enlarged. The cheek teeth are smaller than even
those of the gracile australopithecines. Apparently H. ha-
bilis had a different way of life than the other hominids of
this time.
Cut marks on bones that could have been made by
stone akes have been found at many
sites throughout eastern Africa, dat-
ing from 2 MYA. H. habilis could have
made and used tools in order to strip
meat off these bones and, in keeping
with the size of the teeth, could have
eaten meat to satisfy protein de-
mands. As a scavenger, H. habilis may
have depended simply on the kills of
other animals; or as a predator, he
may have killed small- to medium-
sized prey. This new way of life be-
came available to hominids when
they had the ability to make and use
tools intelligently.
The stone tools made by H. habilis
are called Oldowan tools because they
were rst identied as tools by the
Leakeys at a place called Olduvai
Gorge. Oldowan tools are simple and
look rather clumsy, but perhaps this
hominid also used stone akes. The
akes would have been sharp and
able to scrape away hide and cut ten-
dons to easily remove meat from a
carcass.
H. habilis most likely still ate
fruits, berries, seeds, and other plant
materials. Perhaps a division of
labor arose, with certain members of
a group serving as hunters and oth-
ers as gatherers. Speech would have
facilitated their cooperative efforts,
and later they most likely shared
their food and ate together. In this
way, society and culture could have
begun. Culture, which encompasses
human behavior and products (such as technology and
the arts), is dependent upon the capacity to learn and
transmit knowledge through the ability to speak and
think abstractly.
Prior to the development of culture, adaptation to the
6
5
4
3
2
1
Pleistocene
Holocene
Pliocene
common ancestor
Molecular data puts
ancestor at about 6 MYA.
Relationship to other
hominids is uncertain.
*
Australopithecus
ramidus
Lucy (400 cc)
Australopithecus
afarensis
Australopithecus
africanus
(500 cc)*
Australopithecus
robustus
(500 cc)
Australo-
pithecus
aethiopicus
Australo-
pithecus
boisei
Homo habilis
(700 cc)
Homo erectus
(800
1,200 cc)
M
i
l
l
i
o
n
s

o
f

Y
e
a
r
s

A
g
o

(
M
Y
A
)
Epoch
Modern African
Apes
Homo sapiens
(brain size 1,360 cc)
Cro-Magnon
(above
1,300 cc)
Extinction
environment necessitated a biological change. The acquisi-
tion of culture provided an additional way by which adap-
tation was possible. And the possession of culture by H.
habilis may have hastened the extinction of the australo-
pithecines.
H. habilis warrants classication as a Homo
because of brain size, posture, and dentition.
Circumstantial evidence suggests the use of tools
to prepare meat and also the development of
culture.
Figure 31.16 Recently constructed hominid evolutionary tree.
African apes and hominids split about 6 million years ago (MYA). A. ramidus is the oldest known
of the hominids; A. afarensis lasted about a million years before branching speciation occurred. In
particular, there were gracile and robust australopithecine forms and also the rst human form,
Homo habilis.
Homo erectus
Homo erectus is the name assigned to hominid fossils
found in Asia, and dated between 1.9 and 0.5 MYA. Com-
pared to H. habilis, H. erectus had a larger brain (about 1,000
cc), more pronounced brow ridges, a atter face, and a nose
that projects like ours. These hominids not only stood
erect, they most likely had a striding gait like ours.
It is believed by some that H. erectus rst appeared in
Africa and then migrated into Asia and Europe. Such an ex-
tensive population movement is a rst in the history of hu-
mankind and a tribute to the intellectual and physical skills of
the species. H. erectus was also the rst hominid to use re, and
it fashioned more advanced tools called Acheulean tools,
named after a site in France (Fig. 31.17). There are heavy
teardrop-shaped axes and cleavers as well as akes, which
were probably used for cutting and scraping. Some believe
that H. erectus was a systematic hunter and brought kills to the
same site over and over again. In one location there are over
40,000 bones and 2,647 stones. These sites could have been
home bases where social interaction occurred and a pro-
longed childhood allowed time for much learning. Perhaps a
language evolved and a culture more like our own developed.
H. erectus, which evolved from H. habilis, had a
striding gait, made well-fashioned tools (perhaps
for hunting), and could control re. This hominid
migrated into Europe and Asia from Africa about
1 MYA.
Cro-Magnon
high forehead
flat face
small brow ridge
large brain case
brain size: 1,300 cc
projecting chin
Homo erectus
higher forehead
face almost flat
large brow ridge
large brain case
brain size: 8001,200 cc
smaller zygomatic arch
Homo habilis
forehead present
projecting face
large brow ridge
larger brain case
brain size: 700 cc
large zygomatic arch
Australopithecus
low forehead
projecting face
large brow ridge
small brain case
brain size: 400 cc
large zygomatic arch
Acheulean Oldowan Aurignacian and late Paleolithic
zygomatic
arch
Figure 31.17 Hominid skull anatomy and tools.
The listings compare hominid skulls. Oldowan tools are crude. Acheulean tools are better made and more varied, and Aurignacian tools are well
designed for specic purposes.
Origin of Modern Humans
It is generally recognized that modern humans originated
from H. erectus or a closely related species, but where did
this occur? About 300,000 years B.P. (before present), so-
called archaic Homo sapiens had evolved in Europe, Asia,
and Africa. (Neanderthal is a well-known archaic H. sapiens.)
The hypothesis that each of these individual populations
(with some interbreeding) went on to evolve into modern
humans is called the multiregional continuity hypothesis (Fig.
31.18a). This hypothesis, which proposes no migrations, re-
quires that evolution be essentially similar in several differ-
ent places. Each region would show a continuity of its own
anatomical characteristics from about a million years ago,
when H. erectus rst arrived in Eurasia. Opponents argue
that it seems highly unlikely that evolution would have pro-
duced essentially the same result in these different places.
They suggest, instead, the out-of-Africa hypothesis, which pro-
poses that archaic H. sapiens became fully modern only in
Africa, and thereafter they migrated to Europe and Asia
about 100,000 years B.P. Modern humans may have interbred
to a degree with archaic populations, but in effect, they sup-
planted them (Fig. 31.18b).
Investigators are currently testing two hypotheses:
that modern humans (1) evolved separately in
Europe, Africa, and Asia, and (2) evolved in Africa
and then migrated.
1 MYA
0.5 MYA
Present
Asia Africa Europe
interbreeding
interbreeding
Asia Africa Europe
b. Out of Africa a. Multiregional continuity
Homo erectus evolves into modern humans
in Asia, Africa, and Europe.
migration of
Homo erectus
migration of
Homo erectus
1 MYA
0.5 MYA
Present
Modern humans migrate out of
Africa and replace archaic sapiens
in Asia and Europe. This second
migration occurs about 100,000
years before present (BP).
migration of
Homo erectus
migration of
Homo erectus
Figure 31.18 Origin of modern humans.
a. The multiregional continuity hypothesis proposes that modern humans evolved separately in at least three different places: Asia, Africa, and
Europe. Therefore, continuity of genotypes and phenotypes is expected in these regions. b. The out-of-Africa hypothesis proposes that modern
humans originated only in Africa; then they migrated and supplanted populations of Homo in Asia and Europe about 100,000 years ago.
Homo sapiens
A brain capacity larger than 1,000 cc allows a Homo species
to be classied as Homo sapiens. The Neanderthals (H. sapi-
ens neanderthalensis) take their name from Germanys Nean-
der Valley, where one of the rst Neanderthal skeletons,
dated some 200,000 years old, was discovered. The Nean-
derthals had massive brow ridges, and the nose, the jaws,
and the teeth protruded far forward. The forehead was low
and sloping, and the lower jaw sloped back without a chin.
At this time, the Neanderthals are thought to be an ar-
chaic H. sapiens and most likely not in the main line of Homo
descent. Surprisingly, however, the Neanderthal brain was,
on the average, slightly larger than that of modern humans
(1,400 cc, compared to 1,360 cc in most modern humans).
The Neanderthals were heavily muscled, especially in the
shoulders and the neck. The bones of the limbs were shorter
and thicker than those of modern humans. It is hypothe-
sized that a larger brain than that of modern humans was re-
quired to control the extra musculature. They lived in
Eurasia during the last Ice Age, and their sturdy build could
have helped conserve heat.
The Neanderthals give evidence of being culturally ad-
vanced. They most likely successfully hunted bears, woolly
mammoths, rhinoceroses, reindeer, and other contemporary
animals. They used and could control re, and they even
buried their dead with owers and tools, indicating that
they may have had a religion.
In keeping with the out-of-Africa hypothesis, it is in-
creasingly believed that modern humans, by custom called
Cro-Magnon after a fossil location in France, entered Eura-
sia 100,000 years ago or even earlier. Cro-Magnons
(H. sapiens sapiens) had a thoroughly modern appearance
(Fig. 31.19). They made advanced stone tools called Auri-
gnacian tools (see Fig. 31.17). They were such accomplished
hunters that some researchers believe they were responsible
for the extinction of many larger mammals, such as the giant
sloth, the mammoth, the saber-toothed tiger, and the giant
ox during the late Pleistocene epoch.
Cro-Magnons hunted cooperatively and most likely
lived in small groups, with the men hunting by day while
the women were gatherers and took care of the children. The
Cro-Magnon culture included art. They sculpted small g-
urines out of reindeer bones and antlers. They also painted
beautiful drawings of animals on cave walls in Spain and
France.
The main line of hominid descent is now believed
to include A. ramidus, A. afarensis, H. habilis,
H. erectus, and Cro-Magnon.
Human beings are diverse, but even so, we are all classi-
ed as H. sapiens sapiens. This is consistent with the biologi-
cal denition of species because it is possible for all types of
humans to interbreed and to bear fertile offspring. While it
may appear that there are various races, molecular data
show that the DNA base sequence varies as much between
individuals of the same ethnicity as between individuals of
different ethnicity.
Figure 31.19 Cro-Magnon people.
Cro-Magnon people are the rst to be designated Homo sapiens sapiens. Their toolmaking ability and other cultural attributes, such as their
artistic talents, are legendary.
Summarizing the Concepts
31.1 Echinoderms
Both echinoderms and chordates are deuterostomes. In deuterostomes,
the second embryonic opening becomes the mouth, the coelom forms
by outpocketing of the primitive gut (they are enterocoelomates), and
the dipleurula larva is found among some. Echinoderms (e.g., sea stars,
sea urchins, sea cucumbers, sea lilies) have radial symmetry as adults
(not as larvae) and spines from endoskeletal plates. Typical of echino-
derms, sea stars have tiny skin gills, a central nerve ring with branches,
and a water vascular system for locomotion. Each arm of a sea star con-
tains branches from the nervous, digestive, and reproductive systems.
31.2 Chordates
Chordates (tunicates, lancelets, and vertebrates) have a notochord, a
dorsal hollow nerve cord, pharyngeal pouches, and post-anal tail at
one time in their life history. Tunicates and lancelets are the inverte-
brate chordates. Adult tunicates lack chordate characteristics except
gill slits, but adult lancelets have the three chordate characteristics.
31.3 Vertebrates
Vertebrates have the three chordate characteristics as embryos but the
notochord is replaced by the vertebral column, a part of their en-
doskeleton. The internal organs are well developed and cephalization
is apparent. The vertebrate classes trace their evolutionary history. The
rst vertebrates, represented by hagshes and lampreys, lacked jaws
and ns. Bony shes have jaws and two pairs of ns; the bony shes in-
clude those that are ray-nned and a few that are lobe-nned. Some of
the lobed-nned shes have lungs.
Amphibians evolved from the lobe-nned shes and have two
pairs of limbs. They are represented today by frogs and salamanders.
Frogs usually return to the water to reproduce; frog tadpoles metamor-
phose into terrestrial adults.
Reptiles (todays snakes, lizards, turtles, and crocodiles) lay a
shelled egg, which contains extraembryonic membranes, including an
amnion that allows them to reproduce on land.
Birds are feathered, which helps them maintain a constant body
temperature. They are adapted for ight; their bones are hollow with
air sacs that allow one-way ventilation; their breastbone is keeled, and
they have well-developed sense organs.
Mammals have hair and mammary glands. The former helps
them maintain a constant body temperature, and the latter allow them
to nurse their young. Monotremes lay eggs; marsupials have a pouch
in which the newborn matures, and placental mammals, which are far
more varied and numerous, retain offspring inside the uterus until
birth.
Primates are mammals adapted to living in trees. During the evolution
of primates, various groups diverged in a particular sequence from the
human line of descent. Prosimians (tarsiers and lemurs) diverged rst;
then the monkeys, then the apes. Molecular biologists tell us we are
most closely related to the African apes, whose ancestry split from ours
about 6 MYA.
Human evolution continued in eastern Africa with the evolution
of the australopithecines. The most famous australopithecine is Lucy
(3.4 MYA) whose brain was small but who walked bipedally. Homo ha-
bilis, present about 2 MYA, is certain to have made tools. Homo erectus,
with a brain capacity of 1,000 cc and a striding gate, was the rst to mi-
grate out of Africa.
Two contradicting hypotheses have been suggested about the
origination of modern humans. The multiregional continuity hypothe-
sis says that modern humans originated separately in Asia, Europe,
and Africa. The out-of-Africa hypothesis says that modern humans
originated in Africa and, after migrating into Europe and Asia, re-
placed the archaic Homo species found there. Cro-Magnon is a name of-
ten given to modern humans.
S
ome people believe that animals
should be protected in every way, and
should not be used in laboratory research.
In our society as a whole, the trend is to-
ward a growing recognition of what is
generally referred to as animal rights. In
1985, 63% of Americans polled agreed that
scientists should be allowed to do re-
search that causes pain and injury to ani-
mals like dogs and chimpanzees if it
produces new information about human
health problems. That dropped to 53% in
1995. Psychologists with Ph.D.s earned in
the 1990s are half as likely to express
strong support for animal research as
those who earned their Ph.D.s before 1970.
Those who approve of laboratory re-
search involving animals give examples to
show that even today it would be difcult
to develop new vaccines and medicines
against infectious diseases, develop new
surgical techniques for saving human
lives, or develop treatments for spinal cord
injuries without the use of animals. Even
so, most scientists today are in favor of
what is now called the three Rs: replace-
ment of animals by in vitro, or test-tube,
methods whenever possible; reduction of
the numbers of animals used in experi-
ments; and renement of experiments to
cause less suffering to animals. In the
Netherlands, every scientist starting re-
search that involves animals is well
trained in the three Rs. After designing an
experiment that uses animals, they are
asked to nd ways to answer the same
questions without using animals.
F. Barbara Orlans of the Kennedy In-
stitute of Ethics at Georgetown University
says It is possible to be both pro research
and pro reform. She feels that animal ac-
tivists need to accept that sometimes ani-
mal research is benecial to humans, and
all scientists need to consider the ethical
dilemmas that arise when animals are
used for laboratory research.
Questions
1. Are you opposed to the use of animals in
laboratory experiments? under certain
circumstances? or completely in favor?
Explain.
2. Do you feel that it would be possible for
animal activists and scientists to nd a
compromise they could both accept?
Discuss.
Go To Student OLC
Studying the Concepts
1. What are the general characteristics of deuterostomes? 644
2. What are the general characteristics of echinoderms? Explain
how the water vascular system works in the sea star. 64445
3. What three characteristics do all chordates have at some time
in their life history? Describe the two groups of invertebrate
chordates. 646, 648
4. Discuss the distinguishing characteristics of vertebrates and cite
the milestones in the evolutionary history of vertebrates. 649
5. Describe the three classes of shes. Which have jaws? two
pairs of ns? lungs? 65051
6. Explain in what way amphibians are adapted to life on land
and ways in which they are not adapted. Describe the meta-
morphosis of frogs. 651
7. What is the signicance of a shelled egg? Explain in what
ways reptiles are adapted to life on land. 653
8. What is the signicance of wings? In what other ways are
birds adapted to ying? 655
9. What is the signicance of a constant internal temperature?
What are the three groups of mammals, and what are their
primary characteristics? 65657
10. Name several primate characteristics still retained by
humans. 659
11. Draw and discuss the evolutionary tree for hominids. 661
12. Contrast the multiregional continuity hypothesis and the
out-of-Africa hypothesis. 663
Testing Yourself
Choose the best answer for each question.
1. The tube feet of echinoderms
a. are their head.
b. are a part of the water vascular system.
c. are found in the coelom.
d. help pass sperm to females during reproduction.
e. All of these are correct.
2. Which of these is not a chordate characteristic?
a. dorsal supporting rod, the notochord
b. dorsal hollow nerve cord
c. pharyngeal pouches
d. post-anal tail
e. vertebral column
3. Sharks and bony shes are different in that only
a. bony shes have paired ns.
b. bony shes have a keen sense of smell.
c. bony shes have an operculum.
d. sharks have a bony skeleton.
e. sharks are predaceous.
4. Amphibians arose from
a. tunicates and lancelets. d. ray-nned shes.
b. cartilaginous shes. e. bony shes with lungs.
c. jawless shes.
5. Which of these is not a feature of amphibians?
a. dry skin that resists desiccation
b. metamorphosis from a swimming form to a land form
c. small lungs and a supplemental way of gas exchange
d. reproduction in the water
e. three-chambered heart
6. Which of these is not a difference between reptiles and birds?
ReptilesBirds
a. hard-shelled eggpartial internal development
b. scalesfeathers
c. tetrapodswings
d. cold bloodedwarm blooded
e. no air sacsair sacs
7. Which of these is a true statement?
a. In all mammals, offspring develop within the female.
b. All mammals have hair and mammary glands.
c. All mammals have one birth at a time.
d. All mammals are land-dwelling forms.
e. All of these are true.
8. The most likely line of descent for humans is
a. Common ancestor with an African ape, (a hominoid), Lucy (a
hominid), Homo habilis (a human), Homo erectus, Homo sapiens.
b. prosimians, New World monkeys, Old World monkeys,
Asian apes, African apes, hominids.
c. chimpanzees, orangutan, Australopithecus africanus, marsu-
pials, placental mammals, primates, humans.
d. Only a and b are correct.
e. All of these are correct.
9. Lucy is a member of what species?
a. Homo erectus d. H. habilis
b. Australopithecus afarensis e. A. robustus
c. Neanderthal
10. If the multiregional continuity hypothesis is correct, then
a. hominid fossils in China after 100,000 B.P. are not expected
to resemble earlier fossils.
b. hominid fossils in China after 100,000 B.P. are expected to
resemble earlier fossils.
c. humans evolved in Africa.
d. humans evolved in Eurasia.
e. Both b and d are correct.
11. Which of these is incorrectly matched?
a. H. erectusmade tools
b. Neanderthalgood hunter
c. H. habiliscontrolled re
d. Cro-Magnongood artist
e. A. afarensiswalked erect
12. Complete the following diagram.
c.
d.
a.
b.
Understanding the Terms
amniote egg 649
amphibian 651
anthropoid 659
australopithecines 660
bird 655
bony sh 650
cartilaginous sh 650
chordate 646
Cro-Magnon 664
deuterostome 644
echinoderm 644
feather 655
n 649
gill 646
hominid 660
Homo erectus 662
Homo habilis 661
jaw 650
jawless shes 650
lancelet 648
lung 651
mammal 656
marsupial 657
monotreme 656
Neanderthal 664
notochord 646
placental mammal 657
primate 659
prosimian 659
reptile 653
tunicate 648
vertebrate 649
water vascular system 645
Match the terms to these denitions:
a. Series of canals that takes water to the tube feet of
an echinoderm, allowing them to expand.
b. Group of primates that includes lemurs and tar-
siers and may resemble the rst primates to have evolved.
c. Dorsal supporting rod that exists in all chordates
sometime in their life history; replaced by the vertebral col-
umn in vertebrates.
d. Group of coelomate animals in which rst embry-
onic opening is associated with the anus, and the second
embryonic opening is associated with the mouth.
e. Member of a family that contains humans, and
their direct ancestors which are known only from the fossil
record.
Further Readings for Part 6
Agnew, N., and Demas, M. September 1998. Preserving the Laetoli
footprints. Scientic American 279(3):44. This article recaps the
discovery of hominid footprints in East Africa, and explains
steps taken to preserve them.
Beckage, N. E. November 1997. The parasitic wasps secret
weapon. Scientic American 277(5):82. Some parasitic wasps
produce a virus that suppresses a living hosts immune system.
Ben-Jacob, E., and Levine, H. October 1998. The artistry of
microorganisms. Scientic American 279(4):82. Colonies of
bacteria form geometric patterns, which reect survival
strategies.
Berger, L. August 1998. The dawn of humans: Redrawing our
family tree? National Geographic 194(2):90. South African fossils
show that A. africanus was more apelike than Lucy.
Castro, P., and Huber, M. 1997. Marine biology. 2d ed. St. Louis:
Mosby-Year Book, Inc. This introductory text is designed to
provide a stimulating overview of marine biology.
Chadwick, D. H. March 1998. Planet of the beetles. National
Geographic 193(3):100. With diverse sizes, forms, and functions,
beetles make up one-third of the worlds identied insects.
Chiappe, L. M. September 1998. Wings over Spain. Natural History
107(7):30. The presence of a rst digit on fossil remains of a 115-
million-year-old bird shows advanced ying ability.
Diamond, J. September 1998. Evolving backward. Discover
19(9):64. Studies of the blind mole rat shows how evolved traits
are lost if they are not used.
Erwin, D. E. July 1996. The mother of mass extinction. Scientic
American 275(1):72. Global sea level decline and volcanic
eruptions may have caused mass extinctions.
Foster, K. R., et al. August 1998. The Philadelphia yellow fever
epidemic of 1793. Scientic American 279(2):88. The history of
this epidemic and the possibility of its recurrence are discussed.
Genthe, H. August 1998. The incredible sponge. Smithsonian
29(5):50. Sponge biology and the therapeutic uses of sponges in
treating cancer are discussed.
Gore, R. January 1996. Neanderthals. National Geographic 189(1):2.
Archeological nds are providing much information on the
Neanderthal culture.
. September 1997. The dawn of humans. National
Geographic 192(3):92. Afootprint dated about 117,000 years
ago was discovered in southern Africa.
Thinking Scientically
1. Humans use negative pressure to ll the lungs, and frogs use
positive pressure.
a. What is the difference between negative and positive pres-
sure?
b. Birds have a ow-through systemthe air ows from the
lungs into air sacs and then out by way of a separate set of
tubes. In what way is this more efcient than human
breathing?
c. Frogs have thin, moist skin, and reptiles have thick skin.
Which animal do you predict practices skin breathing?
Which animal most likely has better developed lungs?
d. Humans have a diaphragm, reptiles do not. In what way
does a diaphragm assist breathing?
2. Refer to Figure 31.16 to answer the following questions.
a. What might have caused the evolution of the upright pos-
ture of A. afarensis?
b. How does the hominid evolutionary tree illustrate adap-
tive radiation?
c. How does the evolution of Homo erectus and Cro-Magnon
support the concept of punctuated equilibrium (page 567)?
d. How do you know that all human races should be consid-
ered the same species?
Using Technology
Your study of animals and human evolution is supported by
these available technologies:
Essential Study Partner CD-ROM
Evolution & Diversity Vertebrates
Human Evolution
Visit the Mader web site for related ESP activities.
Exploring the Internet
The Mader Home Page provides resources and tools as
you study this chapter.
http://www.mhhe.com/biosci/genbio/mader
Gwynne, D. T. August 1997. Glandular gifts. Scientic American
277(2):66. Male insects offer body parts and secretions as a
strategy for fertilizing the females eggs.
Johanson, D. C. March 1996. Face-to-face with Lucys family.
National Geographic 189(3):96. New fossils from Ethiopia provide
more information about human evolution.
Kellert, S. R. 1996. The value of life: Biological diversity and human
society. Washington, D.C.: Island Press/Shearwater Books.
Explores the importance of biological diversity to the well-
being of humanity.
Knols, B. G., and Meijerink, J. September/October 1997. Odors
inuence mosquito behavior. Science & Medicine 4(5):56.
Denition of odors may lead to the control of insects that
transmit infectious diseases.
Leakey, M., and Walker, A. June 1997. Early hominid fossils from
Africa. Scientic American 276(6):74. Abone unearthed in 1965
recently proved the existence of a new species of Australo-
pithecus, showing ancestral humans existed 4 million years ago.
Levetin, E., and McMahon, K. 1996. Plants and society. Dubuque,
Iowa: Wm. C. Brown Publishers. Basic botany and the impact of
plants on society are topics covered in this introductory text.
Levy, S. B. March 1998. The challenge of antibiotic resistance.
Scientic American 278(3):46. Misuse and overuse of antibiotics
must end in order to preserve their effectiveness.
Lewin, R. 1997. Patterns in evolution: The new molecular view. New
York: Scientic American Library. This book explores how
genetic information provides insights into evolutionary events.
Lim, D. 1998. Microbiology. 2d ed. Dubuque, Iowa:
WCB/McGraw-Hill. This introductory text shows how
microorganisms relate to one another and to other organisms;
new material in evolution and biodiversity is included.
Line, L. October/November 1998. Fast decline of slow species.
National Wildlife 36(6):22. Box turtles are declining in numbers,
mainly from habitat loss and over-collecting.
Losick, R., and Kaiser, D. February 1997. Why and how bacteria
communicate. Scientic American 276(2):68. Bacteria send and
receive chemical messages and can organize into structures.
Luoma, J. R. March 1997. The magic of paper. National Geographic
191(3):88. The papermaking process is discussed in this article.
Madigan, M. T., and Marrs, B. L. April 1997. Extremophiles.
Scientic American 276(4):86. Certain microorganisms can
withstand extreme environments.
Margulis, L., et al. 1998. Five kingdoms: An illustrated guide to the
phyla of life on earth. 3d ed. New York: W. H. Freeman &
Company. Introduces the kingdoms of organisms.
Martini, F. H. October 1998. Secrets of the slime hag. Scientic
American 279(4):70. The roles of hagsh in ocean ecosystems.
Monastersky, R. March 1998. The rise of life on earth. National
Geographic 193(3):54. Article discusses the origins of microbial
life, stromatolite reefs, and Stanley Millers model of the
primitive atmosphere.
Moore, R., Clark, W. D., et al. 1998. Botany. 2d ed. Dubuque, Iowa:
WCB/McGraw-Hill. This introductory botany text stresses the
importance of plants and the process of science.
Moore-Landecker, E. 1996. Fundamentals of the fungi. 4th ed. Upper
Saddle River, N.J.: Prentice-Hall. For intermediate students, this
text presents a broad introduction to the eld of mycology.
Murawski, D. A. March 1997. Moths come to light. National
Geographic 191(3):40. Moths display a variety of disguises and
survival techniques.
Nester, E. W., Roberts, C. E., et al. 1998. Microbiology: Ahuman
perspective. 2d ed. Dubuque, Iowa: WCB/McGraw-Hill. This
introductory text relates basic microbiology to human health.
Northington, D., and Goodin, J. R. 1996. The botanical world. 2d ed.
St. Louis: Times-Mirror/Mosby College Publishing. This is an
account of plant interactions and basic physiology.
Padian, K., and Chiappe, L. M. February 1998. The origin of birds
and their ight. Scientic American 278(2):38. Recent fossil
discoveries conrm that birds descended from dinosaurs.
Paolella, P. 1998. Introduction to molecular biology. Dubuque, Iowa:
WCB/McGraw-Hill. This introductory text explores the
processes and mechanisms of gene function and control.
Part, M. October 1998. Antarctic desert. National Geographic 4:120.
Microscopic organisms that survive in the Antarctic desert have
been discovered.
Schmidt, G. D., and Roberts, L. S. 1996. Foundations of parasitology.
5th ed. Dubuque, Iowa: Wm. C. Brown Publishers. For upper-
division parasitology classes, this text emphasizes the major
parasites of humans and animals.
Seymour, R. S. March 1997. Plants that warm themselves. Scientic
American 276(3):104. Some plants generate heat to keep
blossoms at a constant temperature.
Sharpe, G. W., et al. 1995. Introduction to forests and renewable
resources. 6th ed. New York: McGraw-Hill, Inc. For forestry
students, this text presents policies and practices in forest
conservation and management.
Shreeve, J. December 1997. Uncovering Patagonias lost world.
December 1997. National Geographic 192(6):120. Recent fossil
nds cause scientists to rethink the evolution of dinosaurs.
Stern, K. 1997. Introductory plant biology. 7th ed. Dubuque, Iowa:
Wm. C. Brown Publishers. This text presents basic botany in a
clear, informative manner.
Sumich, J. L. 1996. An introduction to the biology of marine life. 6th
ed. Dubuque, Iowa: Wm. C. Brown Publishers. This
introductory text covers taxonomy, evolution, ecology, behavior,
and physiology of selected groups of marine organisms.
Sze, P. 1998. Abiology of the algae. 3d ed. Dubuque, Iowa:
WCB/McGraw-Hill. This concise text introduces algae
morphology, evolution, and ecology to the botany major.
Tattersall, I. April 1997. Out of Africa again . . . and again?
Scientic American 276(4):60. Hominids may have migrated out
of Africa several times, with each emigration sending a different
species.
Walters, D. R., and Keil, D. J. 1996. Vascular plant taxonomy. 4th ed.
Dubuque, Iowa: Kendall/Hunt Publishing. Introduces plant
families and experimental aspects of taxonomy.
Webster, D. July 1996. Dinosaurs of the Gobi. National Geographic
190(1):70. Dinosaur fossils are unearthed in the Gobi desert.
Wenke, R. 1996. Patterns in prehistory: Humankinds rst three million
years. 4th ed. New York: Oxford University Press. Provides a
comprehensive review of world prehistory.
Zimmer, C. September 1998. The slime alternative. Discover
19(9):86. Occasionally, individual Dictyostelium amoebas join
together and behave like multicellular organisms.
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