PHYLUM CHORDATA

The phylum Chordata contains three subphyla: two lineages of invertebrates

(Cephalochordata and Urochordata) and the diverse lineage of vertebrates, Vertebrata.
Although the phylum Chordata (kor-dat’ah) (L. chorda, cord) does not have an inordinately
large number of species (about 45,000), its members have been very successful at adapting to
aquatic and terrestrial environments throughout the world.
The phylum is named after the notochord (Gr. noton, the back + L. chorda, cord), a
supportive rod that extends most of the length of the animal dorsal to the body cavity and into
the tail.
Key Morphological Innovations Distinguish Chordates from Other Deuterostome Phyla
Species in the phylum Chordata are distinguished from other deuterostomes by a set of key
morphological innovations: presence of a notochord, post-anal tail, a dorsal hollow nerve
chord, and a pharyngeal slit.
These structures foster higher levels of activity, unique modes of aquatic locomotion, and
more efficient feeding and oxygen acquisition.
NOTOCHORD Early in chordate development, mesoderm that is dorsal to the developing
digestive system forms a notochord (noton = the back; chorda = string). This flexible rod,
constructed of fluid-filled cells surrounded by tough connective tissue, supports the embryo
from head to tail. The notochord forms the skeleton of adult invertebrate chordates. Body
wall muscles are anchored to the notochord, and when these muscles contract, the notochord
bends, but it does not shorten. As a result, the chordate body swings left and right during
locomotion, propelling the animal forward; unlike annelids and other non-chordate
invertebrates, the chordate body does not shorten when the animal is moving. Remnants of
the notochord persist as gelatinous disks in the backbones of adult vertebrates.
SEGMENTAL BODY WALL AND TAIL MUSCLES Chordates evolved in water, and
they swim by contracting segmentally arranged blocks of muscles in the body wall and tail.
The chordate tail, which is posterior to the anus, provides much of the propulsion in aquatic
species. Segmentation allows each muscle block to contract independently; waves of
contractions pass down one side of the animal and then down the other, sweeping the body
and tail back and forth in a smooth and continuous movement.
DORSAL HOLLOW NERVE CORD The central nervous system of chordates is a hollow
nerve cord on the dorsal side of the animal. By contrast, most non-chordate invertebrates
have solid nerve cords on the ventral side. In vertebrates, an anterior enlargement of the nerve
cord forms the brain; in invertebrates, the anterior concentration of nervous system tissue is
described as a ganglion.
PHARYNGEAL SLIT Like the hemichordates described above, most chordates have out
pocketings, perforations, or slits in the pharynx during some stage of the animal’s life cycle.
These paired openings originated as exit holes for water that carried particulate food and
oxygenated water into the mouth. Invertebrate chordates exchange gases with water as it
passes by the walls of the pharynx, acquiring oxygen and eliminating carbon dioxide.
Invertebrate chordates and fishes retain a perforated pharynx throughout their lives. In most

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air-breathing terrestrial vertebrates, the outpocketings or slits are present only during
embryonic development and in larvae.
Members of the subphylum Vertebrata have the following characteristics:
i. Presence of backbone
ii. Gills
iii. Central nervous system
Based on anatomical and physiological traits, the vertebrates belong to the following
traditional groups:
i. Agnatha (Jawless fishes)
ii. Chondrichthyes (Cartilaginous fishes)
iii. Osteichthyes (Bony fishes)
iv. Amphibia (Amphibians)
v. Reptilia (Reptiles)
vi. Aves (Birds)
vii. Mammalia (Mammals)
Agnathans: Hagfishes and Lampreys, Conodonts and Ostracoderms
The earliest vertebrates lacked jaws, but they had a muscular pharynx, which they used to
suck edible titbits into their mouths. Although they do not form a monophyletic group, the
four groups of jawless vertebrates can be described together because their ancestral jawless
condition distinguishes them from the monophyletic Gnathostomata. The two living groups
of agnathans, as well as species that flourished in the Palaeozoic, vary greatly in size and
shape, as well as in the number of vertebrate characters they possess.
Hagfishes and Lampreys Are the Living Descendants of Ancient Agnathan Lineages
Two apparently separate lineages of jawless vertebrates, Myxinoidea (hagfishes) and
Petromyzontoidea (lampreys), still live today. Both have skeletons composed entirely of
cartilage. Although scientists have found no fossilized hagfishes or lampreys from the early
Palaeozoic era, the absence of both jaws and bone in these groups suggests that their lineages
arose early in vertebrate history, before the evolution of bone. Hagfishes and lampreys have a
well-developed notochord, but no true vertebrae or paired fins, and their skin has no scales.
Individuals grow to a maximum length of about 1 m.
The axial skeleton of the 60 living species of hagfishes includes only a cranium and a
notochord; it has no specialized structures surrounding the dorsal nerve cord. Some biologists
do not even include hagfishes among the Vertebrata, because they lack any sign of vertebrae.
Hagfishes are marine scavengers that burrow in sediments on continental shelves. They feed
on invertebrate prey and on dead or dying fishes. In response to predators, they secrete an
immense quantity of sticky, noxious slime; when no longer threatened, a hagfish ties itself
into a knot and wipes the slime from its body. Hagfish life cycles are simple and lack a larval
stage.
The 40 or so living species of lampreys have traces of an axial skeleton. Their notochord is
surrounded by dorsally pointing cartilages that partially cover the nerve cord; many biologists
hypothesize that this arrangement reflects an early stage in the evolution of the vertebral

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column. Most lamprey species are parasitic as adults. They have a circular mouth surrounded
by a sucking disk with which they attach to a fish or other vertebrate host; they feed on a
host’s body fluids after rasping through its skin. In most species, sexually mature adults
migrate from the ocean or a lake to the headwaters of a stream, where they lay eggs and then
die. Their suspension feeding larvae, which resemble adult cephalochordates, burrow into
mud and develop for as long as seven years before undergoing metamorphosis and migrating
to the sea or a lake to live as parasitic adults.

Jawed Fishes
The evolution of jaws is perhaps the greatest of all advances in vertebrate history. Hinged
jaws allow vertebrates to grasp, kill, shred, and crush large food items. Some living species
also use their jaws for defence, for grooming, to construct nests, and to transport their young.

Chondrichthyes Includes Fishes with Cartilaginous Endoskeletons

The 1,050 living species in the Chondrichthyes (chondros = cartilage; ichthys = fish) have
skeletons composed entirely of cartilage, which is much lighter than bone.
Most living chondrichthyans are grouped in the Elasmobranchii, which includes the skates,
rays, and sharks; nearly all are marine predators. Skates and rays are dorsoventrally flattened.
They swim by undulating their enlarged pectoral fins. Most are bottom dwellers that often lie
partly buried in sand. They feed on hard-shelled invertebrates, which they crush with
massive, flattened teeth.
The largest species, the manta ray (Manta birostris), which measures 6 m across, feeds on
plankton in the open ocean. Some rays have electric organs that stun prey with as much as
200 volts.
Sharks are among the ocean’s dominant predators. Flexible fins, lightweight skeletons,
streamlined bodies, and the absence of heavy body armour allow most sharks to pursue prey
rapidly. Their large livers contain copious amounts of oil, which is lighter than water,
increasing their buoyancy. The great white shark (Carcharodon carcharias) is the largest
predatory species, attaining a length of 10 m. The whale shark (Rhincodon typus), which
grows to 18 m, is the largest fish known; it feeds on plankton.
Elasmobranchs (including sharks, skates, and rays) exhibit remarkable adaptations for
acquiring and processing food. Their teeth develop in whorls under the fleshy parts of the
mouth. New teeth migrate forward as old, worn teeth break free. In many sharks, the upper
jaw is loosely attached to the cranium, and it swings down during feeding. As the jaws open,
the mouth spreads widely, sucking in large, hard-to-digest chunks of prey, which are
swallowed intact. Although the elasmobranch digestive system is short, it includes a
corkscrew-shaped spiral valve, which slows the passage of material and increases the surface
area available for digestion and absorption.
Elasmobranchs also have well-developed sensory systems. In addition to vision and olfaction,
they use electroreceptors to detect weak electric currents produced by other animals. And

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their lateral-line system, a row of tiny sensors in canals along both sides of the body, detects
vibrations in water.
Chondrichthyans exhibit numerous reproductive specializations. Males have a pair of organs,
the claspers, on the pelvic fins, which help transfer sperm into the female’s reproductive
tract. Fertilization occurs internally. In many species, females produce large, yolky eggs with
tough, leathery shells. Others retain the eggs within the oviduct until the young hatch. A few
species nourish young within a uterus.
Fishes with Bony Endoskeletons: The Actinopterygii and Sarcopterygii
In terms of diversity and sheer numbers, the Osteichthyes (oste = bone; ichthyes = fishes)—
fishes with bony endoskeletons: a cranium, vertebral column with ribs, and bones supporting
their moveable fins—are the most successful of all vertebrates. The endoskeleton provides
lightweight support, particularly compared with the heavy bony armour of ostracoderms and
placoderms, and enhances their locomotor efficiency. Osteichthyes are diversified into two
lineages. Actinopterygii (aktis = ray; pteron = wing), the ray-finned fishes, have fins that are
supported by thin and flexible bony rays. Sarcopterygii (sarco = flesh), the fleshy-finned
fishes, have fins that are supported by muscles and a stout internal bony skeleton.
Ray-finned fishes have always been more diverse, and they vastly outnumber the fleshy-
finned fishes today. The 28,000 living species of bony fishes occupy nearly every aquatic
habitat and represent more than 95% of living fish species. Adults range from 1 cm to more
than 6 m in length.
Bony fishes have numerous adaptations that increase their swimming efficiency. In many
modern ray-finned fishes, a gas-filled swim bladder serves as a hydrostatic organ that
increases buoyancy. The swim bladder is derived from an ancestral air-breathing lung that
allowed early actinopterygians to gulp air, supplementing their gill respiration in aquatic
habitats where dissolved oxygen concentration was low. The scales of most bony fishes are
small, smooth, and lightweight. And their bodies are covered with a protective coat of mucus,
which retards bacterial growth and smoothen the flow of water.

ACTINOPTERYGII. The most primitive living actinopterygians, sturgeons and

paddlefishes, have mostly cartilaginous skeletons. These large fishes live in rivers and lakes
of the northern hemisphere. Sturgeons feed on detritus and invertebrates; paddlefish consume
plankton. Gars and bowfins are remnants of a more recent radiation. They occur only in the
eastern half of North America, where they feed on fishes and other prey. Gars are protected
from predators by a heavy coat of bony scales.
Teleosts, the latest radiation of Actinopterygii, are the most diverse, successful, and familiar
bony fishes. Evolution has produced a wide range of body forms. Teleosts have an internal
skeleton made almost entirely of bone. On either side of the head, a bony flap of the body
wall, the operculum (plural, opercula), covers a chamber that houses the gills; movements of
the opercula help to bring fresh, oxygenated water across the gills. Sensory systems generally
include large eyes, a lateral-line system, sound receptors, chemoreceptive nostrils, and taste
buds. Variations in jaw structure allow different teleosts to consume plankton, seaweed,
invertebrates, or other vertebrates. Teleosts exhibit remarkable feeding and locomotor

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adaptations. When some teleosts open their mouths, bones at the front of the jaws swing
forward to create a circular opening. Folds of skin extend backward, forming a tube through
which they suck food. Many also have symmetrical caudal fins, posterior to the vertebral
column, which provide power for locomotion. And their pectoral fins lie high on the sides of
the body, providing fine control over swimming. Some species use their pectoral fins for
acquiring food, for courtship, and for care of eggs and young. Some teleosts even use them
for crawling on land or gliding in air.
Most marine species produce small eggs that hatch into planktonic larvae. Eggs of freshwater
teleosts are generally larger and hatch into tiny versions of the adults. In freshwater species,
parents often care for their eggs and young, fanning oxygen-rich water over them, removing
fungal growths, and protecting them from predators. Some freshwater species, such as
guppies, give birth to live young.

SARCOPTERYGII. Two groups of fleshy-finned fishes (Sarcopterygii), the lobe-finned

fishes and lungfishes, are now represented by only eight living species. In the heyday of the
Sarcopterygii, during the Devonian period, their sturdy fins probably allowed them to crawl
or walk on the muddy bottom in the shallow waters they inhabited. Most species probably
also acquired oxygen through primitive lungs to supplement their gill respiration.
Although lobe-finned fishes were once thought to have been extinct for 65 million years, a
living coelacanth (Latimeria chalumnae) was discovered in 1938 near the Comoros Islands,
off the south-eastern coast of Africa. We now know that a population of this meter-long fish
lives at depths of 70 to 600 m, feeding on fishes and squid. Remarkably, a second population
of coelacanths was discovered in 1998, when a specimen was found in an Indonesian fish
market, 10,000 km east of the Comoros population. Based on analyses of its DNA, it is a
distinct species (Latimeria menadoensis).
Lungfishes have changed relatively little over the last 200 million years. Six living species
are distributed on southern continents. The Australian lungfishes, which live in rivers and
pools, use their lungs to supplement gill respiration when dissolved oxygen concentration is
low. The South American and African species, which live in swamps, use their lungs to
collect oxygen during the annual dry season, which they spend encased in a mucus-lined
burrow in the dry mud. When the rains begin, water fills the burrow and the fishes awaken
from dormancy.
Tetrapoda: The Evolution of Limbs
The fossil record suggests that Tetrapoda (tetra = four; pod = foot), the four-limbed
vertebrates and their descendants, arose from a group of fleshy-finned fishes, the
osteolepiforms, in the late Devonian period, about 380 million years ago. Osteolepiforms and
early tetrapods shared several derived characteristics: both had curious infoldings of their
tooth surfaces, a trait with unknown function, and the shapes and positions of bones of the
skulls and appendages were similar.
Key Adaptations Facilitated the Transition to Land
Fishes are not adapted to live on land, and early tetrapods faced serious environmental
challenges as they ventured out of the water. First, because air is less dense than water, it

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provides less support against gravity for an animal’s body. Second, animals exposed to air
inevitably lose body water by evaporation. Third, the sensory systems of fishes, which work
well under water, do not function well in air. However, in swampy, late-Devonian habitats
dry land also offered distinct advantages. Terrestrial plants, soft-bodied invertebrates, and
arthropods provided abundant food, oxygen was more readily available in air than in water,
and predators did not yet live in these new habitats.
In some ways, osteolepiforms had characteristics that may have facilitated the transition to
land. Most had strong, stout fins that enabled them to crawl on the muddy bottom of shallow
pools, and their vertebral column included crescent-shaped bones that provided good support.
They had nostrils leading to sensory pits that housed olfactory (smell) receptors. And they
almost certainly had lungs to augment gill respiration in the swampy, oxygen-poor waters
where they lived.
Over the past few decades, the discovery of new fossils has greatly increased our
understanding of the vertebrates’ transition to land. In 2006, a team of palaeontologists
working in the Canadian arctic discovered the find of a lifetime: Tiktaalik, which in the local
Inuktitut language means “large, freshwater fish.” Tiktaalik, which lived about 375 million
years ago, had characters that identify it as transitional between the osteolepiform fishes and
the tetrapod vertebrates that later colonized the land. Its fishlike characters included bony
scales on its body, fins, and gills, as well as air-breathing lungs.
Its tetrapod characters included a neck; well-developed ribs; a forelimb skeleton that included
a humerus, radius, and ulna; and a flattened skull with upward-pointing eyes. Although
scientists conclude that Tiktaalik was aquatic, it probably lived close to the shore, where it
may have captured prey that lived at the water’s edge. Although the pelvis and pelvic fins
were not described at the time of Tiktaalik’s discovery, a subsequent analysis, published in
2014, revealed that they were larger and more robust than those of the osteolepiforms.
Somewhat later in the Devonian, about 365 million years ago, the earliest tetrapods emerged.
Acanthostega and its close relatives were still somewhat fishlike in the overall form, retaining
a large caudal fin, as well as an operculum; the operculum, which covers the gill chamber in
living fishes, suggests that this animal still used gill respiration.
However, Acanthostega had well-defined digits (that is, fingers and toes)—at least eight on
both the hands and feet. It also had a moderately sturdy vertebral column with small ribs
attached and a well-anchored pelvis. Nevertheless, the structure of its forelimbs suggests that
it could not carry its own weight on land. Thus, paleontologists deduce that, like Tiktaalik, it
occupied shallow-water habitats and captured prey on the shoreline.
In addition to changes in limbs and the vertebral column, life on land also required
adaptations in sensory systems. In fishes, for example, the body wall picks up sound
vibrations and transfers them to sensory receptors directly. But sound waves are harder to
detect in air. Early tetrapods developed a tympanum, a specialized membrane on either side
of the head that is vibrated by airborne sounds. The tympanum connects to the stapes, a bone
that is homologous to the hyomandibula, which had supported the jaws of fishes. The stapes,
in turn, transfers vibrations to the sensory cells of an inner ear.
Modern Amphibians Are Very Different from Their Palaeozoic Ancestors

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Most of the more than 6,130 species of living amphibians (including frogs, salamanders, and
caecelians) are small, and their skeletons contain less bone than those of Palaeozoic tetrapods
such as Acanthostega. All living amphibians are carnivorous as adults, but the aquatic larvae
of some species are herbivores.
Most living amphibians have a thin, scaleless skin, well supplied with blood vessels, that is a
major site of gas exchange. Because gases must enter the body across a thin layer of water,
the skin of most amphibians must remain moist, restricting them to aquatic or wet terrestrial
habitats. Adults of some species also acquire oxygen through saclike lungs. The evolution of
lungs was accompanied by modifications of the heart and circulatory system that increase the
efficiency with which oxygen is delivered to body tissues.
The life cycles of many amphibians (amphi = of both kinds; bios = life) include both larval
and adult stages. Eggs are laid and fertilized in water, where they hatch into larvae, such as
the tadpoles of frogs, which eventually metamorphose into adults. Although the larvae of
many species are aquatic, adults may be aquatic, amphibious, or terrestrial. Some
salamanders are paedomorphic; the larval stage attains sexual maturity without changing its
form or moving to land. By contrast, some frogs and salamanders reproduce on land and skip
the larval stage altogether. But even though they are terrestrial breeders, their eggs dry out
quickly unless they are laid in moist places.
Modern amphibians are represented by three clades: Anura, Urodela and Gymnophiona.
ANURA. The 5,400 species of frogs and toads have short, compact bodies, and adults lack
tails. Their elongate hind legs and webbed feet allow them to hop on land or swim. A few
species are adapted to dry habitats, withstanding periods of drought by encasing themselves
in mucous cocoons.
CAUDATA. Most of the 560 species of salamanders and newts have an elongate, tailed body
and four legs. They walk by alternately contracting muscles on either side of the body much
the way fishes swim. Species in the most diverse group, the lungless salamanders, are fully
terrestrial throughout their lives, using their skin and the lining of the throat for gas exchange.
GYMNOPHIONA. The 170 species of caecelians (Gymnophiona, from gymnos = naked;
ophioneos = snakelike) are legless burrowing amphibians with wormlike bodies. They
occupy tropical habitats throughout the world. Unlike other modern amphibians, caecelians
have small bony scales embedded in their skin. Fertilization is internal, and females give birth
to live young.
Reptiles: Diapsid Amniotes
Class Reptilia. Members of the class Reptilia possess amniotic eggs, which develop free
from standing or flowing water. Numerous other adaptations have allowed members of this
class to flourish on land.
All living reptiles are members of the Diapsida (Gr. di, two). Members of this group have
upper and lower openings in the temporal region of the skull.
Fossil evidence and a broad phylogenomic study reveal that Reptilia differentiated into two
major lineages, the Lepidosauria (lepis = scale; sauros = lizard) and the Archelosauria
(arkhon = ruler; chelon = turtle), which differ in many skeletal and genetic characteristics.

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Living diapsids include the Lepidosauria (snakes, lizards, and tuataras) and Archosauria
(crocodiles).
Reptiles are characterized by a skull with one surface (condyle) for articulation with the first
neck vertebra, respiration by lungs, metanephric kidneys, internal fertilization, and amniotic
eggs. Reptiles also have dry skin with keratinized epidermal scales. Keratin is a resistant
protein found in epidermally derived structures of amniotes. It is protective, and when
chemically bonded to phospholipids, prevents water loss across body surfaces.

Testudines: Turtles
Turtles Have Bodies Encased in a Bony Shell
The turtle body plan, largely defined by a bony, boxlike shell, has changed little since the
group first appeared during the Triassic period, nearly 250 million years ago. The shell
includes a dorsal carapace and a ventral plastron. A turtle’s ribs are fused to the inside of the
carapace, and, in contrast to other tetrapods, the pectoral and pelvic girdles lie within the
ribcage.
Large, keratinized scales cover the bony plates that form the shell. The 330 living species of
turtles occupy terrestrial, freshwater, and marine habitats. They range from 8 cm to 2 m in
length. All species lack teeth, but they use a keratinized beak and powerful jaw muscles to
feed on plants or animal prey. When threatened, most species retract into their shells. Many
species are now highly endangered because adults are hunted for meat, their eggs are
consumed by humans and other predators, and their young are collected for the pet trade.

Crocodilia: Crocodiles
Crocodilians are semiaquatic, predatory archosaurs. The 21 living species of alligators and
crocodiles (Crocodilia, from crocodilus = crocodile) are the remnants of a once-diverse
archosaur lineage. The largest species, the Australian saltwater crocodile (Crocodylus
porosus), grows to 7 m. Crocodilians are aquatic predators that consume other vertebrates.
Striking anatomical adaptations distinguish them from living lepidosaurs, including a four
chambered heart and a one-way flow of air through their lungs, characters that are
homologous to those of birds.
American alligators (Alligator mississippiensis) exhibit strong maternal behaviour, which
also reflects their relationship to birds. Females guard their nests ferociously and help their
young break out of their eggshells. The young stay close to the mother for about a year,
feeding on scraps that fall from her mouth and living under her watchful protection.

Living Lepidosaurs: Sphenodontids and Squamates

Two groups of lepidosaurs (Sphenodontida and Squamata) still live today. In addition to
sharing certain skeletal characteristics, their bodies are encased in a dry, scaly skin.
Squamates—Lizards and Snakes—Are Covered by Overlapping, Keratinized Scales.

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The skin of lizards and snakes (Squamata, squama = scale) is composed of overlapping,
keratinized scales that protect against dehydration. Squamates periodically shed their skin as
they grow, much the way arthropods shed their exoskeletons.
Most squamates regulate their body temperature behaviourally; they are active only when
weather conditions are favourable, and they shuttle between sunny and shady places when
they need to warm up or cool down.
Most of the 6,000 lizard species are less than 15 cm long. However, the Komodo dragon
(Varanus komodoensis) grows to nearly 3 m. Lizards occupy a wide range of habitats, but
they are especially common in deserts and the tropics; one species (Zootoca vivipara) occurs
within the Arctic Circle. Most lizards feed on insects, although some eat leaves or meat.
The 3,400 species of snakes evolved from a lineage of lizards that lost their legs over
evolutionary time. Streamlined bodies make snakes efficient burrowers or climbers.
Many subterranean species are only 10 or 15 cm long, but the giant constrictors may grow to
10 m. Unlike lizards, all snakes are predators that swallow prey whole. Snakes have thinner
skull bones than their lizard ancestors did, and the bones are connected to each other by
elastic ligaments that stretch remarkably, allowing some snakes to swallow food that is larger
than their heads. Snakes also have well-developed sensory systems for detecting prey. The
flicking tongue carries airborne molecules to sensory receptors in the roof of the mouth. Most
snakes can detect vibrations on the ground, and some, such as rattlesnakes, have heat-sensing
organs. Many snakes kill by constriction, which suffocates prey, and several groups produce
toxins that immobilize, kill, and partially digest the prey before it is swallowed.

AVES: Birds
Birds (Aves) appeared in the Jurassic period as descendants of carnivorous, bipedal
dinosaurs. Thus, they are full-fledged members of the archosaur lineage. Their evolutionary
relationship to dinosaurs is evident in their skeletal anatomy, the scales on their legs and feet,
and their posture when walking. However, powered flight gave birds access to new adaptive
zones, setting the stage for their astounding evolutionary success.
Birds Have Key Adaptations That Reduce Body Weight and Power Flight
The skeletons of birds are both lightweight and strong. For example, the endoskeleton of the
frigate bird, which has a 1.5 m wingspan, weighs little more than 100 g, far less than its
feathers. Most birds have hollow limb bones with small supporting struts that crisscross the
internal cavities. Evolution has also reduced the number of separate bony elements in the
wings, skull, and vertebral column (especially the tail), making the skeleton light and rigid.
And all modern birds lack teeth, which are dense and heavy; they acquire food with a
lightweight, keratinized bill. Many species have a long, flexible neck, which allows them to
use their bills for feeding, grooming, nest-building, and social interactions.
The bones associated with flight are generally large. The forelimb and forefoot are elongate,
forming the structural support for the wing. And most modern birds possess a keeled sternum
(breastbone) to which massive flight muscles are attached. Not all birds are strong fliers,
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however; ostriches and other bipedal runners have strong, muscular legs but small wings and
flight muscles.
Like the skeleton, soft internal organs are modified in ways that reduce weight. Most birds
lack a urinary bladder; uric acid paste is eliminated with digestive wastes. Females have only
one ovary and never carry more than one mature egg; eggs are laid as soon as they are
shelled.
All birds also possess feathers, sturdy, lightweight structures derived from scales in the skin
of their ancestors. Each feather has numerous barbs and barbules with tiny hooks and grooves
that maintain the feathers’ structure, even during vigorous activity. Flight feathers on the
wings provide lift, contour feathers streamline the surface of the body, and down feathers
form an insulating cover close to the skin. Worn feathers are moulted and replaced once or
twice each year.
Other adaptations for flight allow birds to harness the energy needed to power their flight
muscles. Their metabolic rates are eight to ten times higher than those of other comparably
sized Reptilia, and they process energy-rich food rapidly. A complex and efficient respiratory
system and four-chambered heart enable them to consume and distribute oxygen efficiently.
As a consequence of high rates of metabolic heat production, birds maintain a high and
constant body temperature.

MAMMALIA: Monotremes, Marsupials, and Placentals

The synapsid lineage, which includes the living mammals, was the first group of amniotes to
diversify broadly on land.
Two mammalian lineages, the egg-laying Prototheria (or monotremes) and the live-bearing
Theria (marsupials and placentals), survived the mass extinction that eliminated most
dinosaurs at the end of the Mesozoic. The Theria diversified into the mammalian groups that
are most
familiar today.
Mammals Exhibit Key Adaptations in Anatomy, Physiology, and Behaviour
Four sets of key adaptations fostered the success of mammals.

HIGH METABOLIC RATE AND BODY TEMPERATURE Like birds, mammals have
high metabolic rates that liberate enough energy from food to maintain high activity levels
and enough heat to maintain high body temperatures. An outer covering of fur and a layer of
subcutaneous fat help retain body heat. Using metabolic heat to stay warm requires lots of
oxygen, and mammals have a muscular organ, the diaphragm, that fills their lungs with air.
Four-chambered hearts and complex circulatory systems deliver oxygen to active tissues.
SPECIALIZATIONS OF THE TEETH AND JAWS Mammals also have anatomical
features that allow them to feed efficiently. Ancestrally, mammals have four types of teeth:
flattened incisors nip and cut food; pointed canines pierce and kill prey; and two sets of

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cheek teeth, premolars and molars, grind and crush food. Moreover, teeth in the upper and
lower jaws occlude (that is, fit together) tightly as the mouth is closed; thus, mammals can
use their large jaw muscles to chew food thoroughly. The occlusion of premolars and molars
allows some mammals to feed on exceptionally tough plant material, such as grasses and
twigs.
PARENTAL CARE Mammals provide substantial parental care to their young. In most
species, young complete development within a female’s uterus, deriving nourishment through
the placenta, a specialized organ that mediates the delivery of oxygen and nutrients. Females
also have mammary glands, specialized structures that produce energy rich milk, a watery
mixture of fats, sugars, proteins, vitamins, and minerals. This perfectly balanced diet is the
sole source of nutrients for new born offspring.
COMPLEX BRAINS Finally, mammals have larger brains than other tetrapods of
equivalent body size; the difference lies primarily in the cerebral cortex, the part of the
forebrain responsible for information processing and learning. Most mammals have a keen
sense of olfaction; as Molecular Insights explains, a huge variety of olfactory receptor
proteins send information to the parts of the cerebral cortex that allow mammals to sense and
interpret a wide variety of odours. Extensive postnatal care provides opportunities for
offspring to learn from older individuals. Thus, mammalian behaviour is strongly influenced
by past experience and learning, as well as by genetically programmed instincts.
The major groups of modern mammals differ in their reproductive adaptations.
Biologists recognize a primary distinction between two lineages of modern mammals: the
egg-laying Prototheria, or monotremes, and the live-bearing Theria. The Theria, in turn,
diversified into two sublineages: the Metatheria, or marsupials, and the Eutheria, or
placentals, which also differ in their reproductive adaptations.

MONOTREMES The five living species in Prototheria (protos = first; therion = wild beast),
the monotremes, which are limited to the Australian region, reproduce with a leathery shelled
egg. Newly hatched young lap up milk secreted by modified sweat glands on the mother’s
belly. The four species of echidnas, or spiny anteaters, feed on ants or termites. The duck-
billed platypus (Ornithorhynchus anatinus) lives in burrows along riverbanks and feeds on
aquatic invertebrates.
MARSUPIALS The 270 species of Metatheria (meta = between), the marsupials, have short
gestation; the young are nourished through a placenta very briefly—sometimes only for 8 to
10 days—before birth. New-borns use their forelimbs to drag themselves across the mother’s
belly fur and enter her abdominal pouch, the marsupium, where they complete development
attached to a teat. Marsupials are the dominant native mammals of Australia and a minor
component of the South American fauna; only one marsupial species, the opossum
(Didelphis virginiana), occurs in North America.
PLACENTALS The 4,000 species of Eutheria (eu = true), the placental mammals, are the
dominant mammals today. They complete embryonic development in the mother’s uterus,
nourished through a placenta until they reach a fairly advanced stage of development. Some
species, such as humans, are helpless at birth, but others, such as horses, are quickly mobile.

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Biologists divide eutherians into about 18 groups, traditionally identified as orders, only eight
of which include more than 50 living species. Rodents (Rodentia) make up about 45% of
eutherian species, and bats (Chiroptera) comprise another 22%. Our own group, Primates, is
represented by fewer than 250 living species (less than 5% of all mammalian species), many
of which are highly endangered.
Some eutherians have highly specialized locomotor structures. For example, whales and
dolphins (Cetacea) are descended from terrestrial ancestors, but their appendages do not
function on land; they are now restricted to aquatic habitats. By contrast, seals and walruses
(Carnivora) feed underwater but rest and breed on land. Bats (Chiroptera) use wings for
powered flight.
Although early mammals appear to have been insectivorous, the diets of modern eutherians
are diverse. Odd-toed ungulates (ungula = hoof) such as horses and rhinoceroses
(Perissodactyla), even-toed ungulates such as cows and camels (Artiodactyla), and rabbits
and hares (Lagomorpha) feed on vegetation. Lions and wolves (Carnivora) consume other
animals. Many hedgehogs, moles, and shrews (Lipotyphla) and bats eat insects, but some
feed on flowers, fruit, and nectar. Many whales and dolphins prey on fishes and other
animals, but some eat plankton. And some groups, including rodents and primates, feed
opportunistically on both plant and animal matter.

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