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Four distinctive derived characteristics of chordates distinguish them from their


A. Notochord, or a rod of vacuolated cells, encased by a firm sheath that

lies ventral to the neural tube in vertebrate embryos and some adults.

B. Hollow nerve cord that lies dorsal to the notochord

C. Pharyngeal pouches

D. Endostyle - elongated groove in the pharynx floor of protochordates

that may develop as the thyroid gland in chordates

In the subphylum Vertebrata, all members possess the

four chordate characteristics at some time in
development, but often these structures are altered
significantly in adult animals. Subphylum
Vertebrates constitute the vast majority of living chordates, and they have evolved an
enormous variety of forms. The backbone of vertebrates protects the nerve cord and
serves as the axis of the internal skeleton. The skeleton provides strength and rigidity to
the body and is an attachment site for muscles. The vertebrae in the middle region of the
trunk give rise to pairs of ribs, which surround and protect the internal organs. A
cartilaginous or bony case encloses the brain. Bone is a substance unique to vertebrates. It
was formerly thought that vertebrates with cartilage skeletons (cyclostomes and sharklike
fishes) were descended from early vertebrates that had not yet developed bone. However,
very primitive fishes with bone skeletons are known from the fossil record, so lack of
bone is now believed to be a degenerate rather than a primitive feature. All but the most
primitive vertebrates, known as jawless fishes, have jaws and paired appendages. The
fishes and, to a lesser extent, the amphibians and reptiles show a segmental arrangement
of the muscles of the body wall and of the nerves leading to them.
There are eight vertebrate classes. Four are aquatic, and may be grouped together as the
superclass Pisces, or fish; four are terrestrial or (in the case of amphibians)
semiterrestrial, and may be grouped as the superclass Tetrapoda, or four-footed animals.
Fishes breathe water by means of gills located in internal passages, although they may
also have lungs as supplementary air-breathing organs. Most move through the water by
weaving movements of the trunk and tail. All have fins, and most have two sets of paired
fins (pelvic and pectoral). Tetrapods breath air, usually by means of lungs, and never
have gills as adults, although the amphibians go through a gilled, water-breathing stage.
Except where the appendages have been lost, as in snakes, all have two pairs of limbs,
generally used for locomotion; these are homologous to the pelvic and pectoral fins of

Class Agnatha

The Agnatha, or jawless fishes, are the oldest known vertebrates. The only surviving
members of this class are the hagfish and lampreys, known as cyclostomes. Cyclostomes
have long, slender bodies with dorsal, ventral, and caudal (tail) fins, all in the median


Although in their lack of jaws or paired

lateral appendages they represent a very primitive stage of vertebrate development, the
modern cyclostomes are highly adapted for their particular ways of life. The hagfish is a
specialized scavenger, and the lamprey is a parasite on other fishes. The lamprey has a
round mouth without skeletal supports, a rasping tongue, and a single, dorsally located
nostril. The gill passages are enlarged to form pouches and are lined with gill filaments
that serve as a surface for the exchange of respiratory gases; in vertebrates the gill
passages have acquired a respiratory function. In cyclostomes, as in all fishes, water is
taken in through the mouth and expelled through the gill passages; as water passes over
the thin-walled gill filaments, dissolved oxygen diffuses into the blood, and carbon
dioxide diffuses out. The lamprey has a notochord extending from the head to near the tip
of the tail. A few cartilaginous blocks around the notochord constitute the bare rudiments
of a backbone; a cartilage framework supports the gill region, and there is a rudimentary
cartilage braincase. The meagerness of the skeleton is considered a degenerate, not a
primitive condition. The larva of the marine lamprey is a small animal, resembling a
lancelet, that uses the pharynx and gill passages for filter-feeding. It metamorphoses into
the adult form before migrating to the sea. The extinct relatives of the cyclostomes, called
ostracoderms, were jawless fishes with bony armor and in some cases a well-developed
bony skeleton.

Class Placodermi

The placoderms, an entirely extinct group of armored fishes, were the first jawed
vertebrates. Jaws enabled vertebrates to become predators, an important factor in the later
development of active, complex forms. The placoderms were also the first vertebrates to
have the two pairs of lateral appendages (supported by pelvic and pectoral girdles) that
characterized all later vertebrate groups. These primitive paired fins gave rise to the
pelvic and pectoral fins of modern fishes and to the limbs of four-footed animals. The
ostracoderms are thought to have given rise to both the sharklike and the bony fishes.

Class Chondrichthyes

The almost exclusively marine sharks, rays, and chimaeras of the class Chondrichthyes
have skeletons made of cartilage. The mouth, equipped in most sharks with numerous
sharp teeth, is located on the underside
of the head. Passages called gill arches lead from the pharynx to the exterior and are lined
with gill filaments. The gill arches are supported by gill bars. Except in chimaeras, the
external gill slits are not covered and are conspicuous on the surface of the body. The jaw
consists of two distinct pieces; the upper part is not fused to the braincase as in higher
vertebrates. The tail is asymmetrical, curving upward in a shape found in early fossil
fishes and thought to be primitive. There is no lung or swim bladder. The skin is studded
with toothlike structures called denticles. Sharks have typical vertebrate kidneys that
excrete a very dilute urine consisting mostly of water; presumably the earliest vertebrates
(ancestral to sharks) evolved in freshwater, where this function is necessary to maintain
the correct concentration of the physiologically important salts in the tissues against the
tendency for them to be diluted by the inward diffusion of water. In marine species, on
the other hand, it is necessary to prevent the concentration of those salts from increasing.
Although the kidneys of sharks pump out water, their body fluids contain ammonia in
concentrations high enough to make the osmotic pressure equal to that of seawater; this
prevents the inward diffusion of salts. Sharks have internal fertilization and lay large
eggs, well supplied with yolk and protected by leathery shells. In a few species the eggs
are hatched within the body.

Class Osteichthyes
The bony fish of the class Osteichthyes are the predominant class of living fishes. In this
group the bony skeleton has been retained and lungs and swim bladders have evolved.
Early bony fishes evolved in freshwater under conditions of periodic drought and
stagnation and developed an internal, moisture-retaining organ, the lung, for gas
exchange. Those fishes gave rise to two lines of descendants.

Members of one line, the fleshy-finned fish, had thick fins with supporting bones, used
for crawling. The only survivors of that group are the coelacanth, or lobefin, which has a
vestigial lung and crawls on the seafloor, and the freshwater lungfishes of drought-ridden
areas, which can crawl over land in search of water and even live out of water for several
years. Early fleshy-finned fish gave rise to the first land vertebrates, the amphibians.

The second line, the ray-finned fish, constitutes the predominant modern group. Ray-
finned fish are highly specialized for aquatic life; they have developed thin, lightweight
fins supported by slender rays, and used only for balance and steering. The lung, a ventral
outpocketing of the pharynx, was no longer necessary as these fish invaded freshwaters
and oceans throughout the world; it shifted to a dorsal position and evolved into a
hydrostatic organ called the swim bladder, or air float. The swim bladder, along with the
strong, lightweight skeletal construction, makes ray-finned fishes much lighter-bodied
than sharks. The gill passages of ray-finned fishes resemble those of sharks, but have a
bony covering, called the operculum, over the external gill slits. Ray-fins have a typical
vertebrate kidney which, in freshwater forms, maintains the proper salt concentration in
the tissues by excreting excess water. In the marine forms the activity of the kidney is
offset by the activity of salt-secreting glands; in addition, the kidney may be modified so
as to produce a more concentrated urine. The heart, like that of sharks, has two chambers,
and there is no separation of oxygenated and deoxygenated blood in the circulatory
system. A few primitive ray-fins (the sturgeon, the paddle fish, and the bowfin) have
asymmetrical tails and thick scales regarded as primitive in construction.

The higher ray-fins, or teleosts, have more or less symmetrical tail fins extending above
and below the vertebral column, and typical fish scales made of very thin layers of bone.
Most marine teleosts produce enormous numbers of small eggs that are externally
fertilized and float in plankton; only a few of these survive. In many species there is a
larval stage that is quite dissimilar to the adult. Teleosts have evolved a tremendous
variety of forms and occupy very diverse ecological niches, both freshwater and marine.

Class Amphibia

The amphibians, the first vertebrates to have limbs, evolved during the Devonian period.
They are only partially terrestrial: Their externally fertilized eggs are laid in freshwater,
and they go through a gilled, aquatic larval stage (the tadpole stage) before
metamorphosing into land-living adults. The skin of the adult is water-permeable, and the
animal must live in a moist environment to prevent desiccation. The adult usually
breathes by means of lungs, although some breathe directly through the skin. The heart is
a three-chambered structure that creates a partial separation between oxygenated blood,
destined for the body tissues, and depleted blood, destined for the lungs; this provides
better oxygenation than does a system in which the two kinds of blood mix. There are
only three groups of amphibians living today. The salamanders are closest to the basic
amphibian stock in form and in method of locomotion. Although supported by limbs,
they move with a wriggling motion similar to that of a fish. The frogs and toads are
specialized for jumping, with long, muscular hind legs, while the tropical caecilians are
burrowing forms that have lost all but vestigial traces of their limbs.

Class Reptilia

The reptiles, which evolved from amphibians during the Carboniferous period, were the
first vertebrate group to become entirely independent of water. This was made possible
by the development of a scaly, water-resistant skin and of the terrestrial, or amniote, type
of egg found in all higher land vertebrates. The amniote egg has an elaborate series of
internal membranes (one of which is called the amnion) surrounding a pool of liquid in
which the embryo develops; the membranes prevent desiccation and allow inward
diffusion of oxygen. Reptilian eggs have porous shells and large amounts of yolk.
Fertilization is internal. In most cases the eggs are laid unhatched; in a few species they
are retained and hatched in the body. Reptiles, including such forms as turtles and sea
snakes that have returned to an aquatic life, are air-breathing at all stages, and nearly all
lay their eggs on land. Gill passages appear, as in birds and mammals, only in the

During the Mesozoic era, reptiles were exceedingly diverse and numerous. The reptilian
dinosaurs included the largest terrestrial animals that have ever lived, as well as many
smaller forms. There were also flying and aquatic reptiles. With the rise of the early
mammals the decline of the reptiles began. The only large and successful modern group
of reptiles is the order of lizards and snakes. Snakes are descended from lizards, but have
lost their limbs. Reptiles, like fish and amphibians, are cold-blooded, that is, they have
little ability to regulate their body temperature, which approaches that of the
environment. The reptiles gave rise to the two warm-blooded vertebrate groups, the birds

and the mammals.

Class Aves

The birds evolved from reptiles in the Jurassic period. Their front limbs are modified into
wings, and the breastbone is greatly enlarged to support flight muscles. They have an
insulating covering of feathers, which has been an important factor in their ability to
regulate body temperature. The other advance that enabled birds to become warm-
blooded was the evolution of a four-chambered heart, making the circulatory system a
complete double circuit: oxygenated blood is pumped from the lungs to the tissues, and
deoxygenated blood is pumped from the tissues to the lungs. The only major group
besides insects to invade the air, birds are much less restricted by external temperature
requirements than cold-blooded animals, and they have spread throughout every part of
the world. They live in many kinds of habitat and have evolved a diversity of forms.
Some have become flightless terrestrial animals, while others are aquatic, using their
wings for swimming instead of or in addition to flying. Fertilization is internal. The eggs
of birds are similar to those of reptiles, but parental care of the eggs and young is highly


Class Mammalia

The mammals also arose from reptiles in the Jurassic period and are now the dominant
form of terrestrial vertebrate life. Like the birds, they have a four-chambered heart and a
double-circuit circulatory system and are able to regulate body temperature. In the case of
mammals the insulating covering is provided by hair, a feature unique to the class,
although in a few forms (particularly in marine species) nearly all the hair is lost, and
insulation is provided by fat. A second distinguishing characteristic of mammals is the
production of milk by the females for the nourishment of the young. All mammals have
internal fertilization, and all but the most primitive (the egg-laying monotremes of
Australia) bear live young. The mammalian egg contains little yolk. In the marsupials the
young are born at an extremely undeveloped stage and continue to develop in a milk-
supplied pouch. In the vastly more numerous placental mammals nourishment is passed
from the circulatory system of the mother to that of the embryo by means of a placenta,
and the young are born well-developed. Most mammals have highly evolved sense
organs and larger brains than other vertebrates. As a group they display great adaptability
to a variety of conditions and have spread to all regions of the world.

The earliest placental mammals were small animals of the insectivore type, but adaptive
radiation has resulted in great diversity of forms and ways of life. Some mammals are
predators; others are herbivores with specialized digestive systems. Some have taken up
an aquatic existence and a few marine forms (whales and sirenians) even give birth at sea.
Members of one group, the bats, have developed membranous wings supported by
elongated fingers and lead an aerial existence. The primates, the group that includes
humans, are fairly close to the original mammalian type in general structure (for example,
they have five fingers and toes and walk flat on the sole of the foot), but they have
undergone great evolutionary advances in the development of the brain, vision, and
manual dexterity.