Fish are incredible diverse group, made up of three living classes that

hold an important place both in modern ecology and in evolutionary

history. Fish are incredibly successful, they are numerous and have a
huge variety of adaptations. They are successful in every water lled
habitat. Furthermore, sh are the rst known vertebrates and also the
stepping stone to all land-walking vertebrates (tetrapods).

What are tetrapods: The superclass Tetrapoda, or the tetrapods,
comprises the rst four-limbed vertebrates and their descendants,
including the living and extinct amphibians, reptiles, birds, and mammals.




Fish are complicated, diverse, and have a long evolutionary history.
In order to understand the place of sh as both a transitory group and as
a modern one it is important to understand the times in which and from
which they evolved. Fish are the rst known true chordates. The rst
vertebrate that has been found is the Upper Cambrian fossil Anaspis,
which is more than 500 million years old (3). This fossil, while being
fragmentary, is thought to be an armored, jawless sh (3). Fish did most of
their evolving between ve million and three and a half million years ago.
These two periods were known as the Silurian and the Devonian periods.
In yhe middle Silurian, the jawless shes had diversied, but it was not
until the Devonian that the true variety of shes really ourished. In fact,
the Devonian, is often referred to as the "Age of Fishes". Towards the end
of the Devonian the rst tetrapods (vertebrates which evolved true legs
with which they could walk on land) had evolved from one specic branch
of sh. Fish greatly specialized in their aquatic niche during both the
Devonian and the Silurian and part of this evolution led to adaptations to
land in the form of amphibians (9).

Fish are in no way simply a stepping stone to amphibians, they are a
much more signicant than this. They have evolved to be masters of their
domain, the water. They come in many forms, have the ability to eat a
huge variety of foods, and have populated almost every body of water. In
fact, sh are the most common vertebrate, with there being approximately
24,000 species alive today. This number is mind boggling when put in
perspective; the next most common vertebrate are birds with a mere
8,600 species (8). This multitude of species ranges in size, morphology,
agility, and adaptations to environment (10). Fish have been broken down
into a series of classes that separate them based on characteristics.
These groups help to classify the wide variety of species that make up
sh and help lead to understand of the current and evolutionary niche of
sh.

The rst sh to evolve were the Agnathans (Class agnatha) (7). These
jawless shes are the rst vertebrates. These sh have round mouth parts
that could be used for sucking or lter feeding (10). These rasping,
sucking mouths are currently found on modern lampreys and hagshes.
These sh were often extremely armored in order to help them protect
themselves. One group that evolved before the Silurian were the
Ostracoderms which have been described as "small, blunt-headed
forms...they fed on debris in the mud, bullet shaped swimmers, and
some...with an astonishing array of spines and crests on their heads." (1).
Most of these types of sh are currently extinct with the exceptions of the
lampreys and the hagsh.

From these bottom feeding, jawless sh came the evolution of jawed sh.
Jaws evolved only once (rather than evolving multiple times in di"erent
species through parallel evolution). Jaws evolved from gill arches which
are the bony parts between gill slits. It is thought that a gill arch in an
agnathan became fused to its skull (11). The upper part of the gill support
became the top jaw and the bottom part of the gill support became the
bottom jaw. Embryology points to this and the arrangement of nerves in
shark heads and most simple shes shows that jaws are in line with gill
arches. While sh had the rst bony jaws, they also have some of the
most complicated. While the human head has only one moving part (the
jaw), the head of a sh may have more than twenty-four bones that may
move together in feeding (8). The evolution of the jaw is incredibly
important because it led to sh to be able to ingest a much wider variety
of foods and allowed them to be active hunters as opposed to passive
lter feeder (1). This led to a wide variety of adaptations in morphology.
Fish became more agile to be better predators, they were able to reduce
their armor because they were less vulnerable, and their muscle density
was able to decrease becuase they no longer led such a sluggish lifestyle
(10).

The rst jawless shes include the acnathodians and placoderms. Even
early in their development the jawless shes showed great diversity. The
acnathodians appeared around four and half million years ago. They were
the rst jawed shes to evolve and they tended to be small, streamlined,
and had huge eyes. Placoderms on the other hand appeared in the early
Silurian and dominated the Devonian. These were some strange looking
sh! As described by Peter Gore, "the Placoderms had bony armor that
covered the head and forepart of the body. In many, a movable joint
between the head and body armor let the head rock back to open the
mouth wide. The primitive jaws had jagged bony edges that served as
teeth. The tail end usually lacked protection"(11). At the beginning of the
Devonian, Placoderms were small, but they soon increased in size. The
Dunkleosteus, a placoderm, grew to be as large as 35 feet, had well
developed jaws, with fang like teeth. The front of the trunk was heavily
armored and the hind part was either bare or covered with small scales
(11). The Dunkleosteus and all the other large placoderms are extinct, but
in the Devonian they dominated both salt and freshwater.

While the huge, menacing placoderms have long since vanished from the
earth, members of the Class Chondritchthyes, some of which are equally
menacing, have not. This class includes the sharks, skates, and rays
(along with some other sh) and these rst evolved between 400 and 450
million years ago (1, 4). They probably shared a common ancestor with
the placoderms (1). This class is commonly refer to as the cartilaginous
sh because they lack true bone, instead they have cartilage and calcied
cartilage for internal support. This type of skeleton is extremely light and
exible and helps these species be agile predators. Chondrithyes have
evolved two kinds of jaws; sharks have extremely powerful biting and
crushing jaws while skates and ray uses their jaws to be bottom feeders
searching for mollusks (4). Sharks are often referred to as living fossils
since they have not done much evolving since the Late Paleozoic. This is
not to say that the sharks that exist now are they only sharks ever to exist.
After the decline of the placoderms, there was a shark explosion that led
to sharks with mouth's lled with coils of serrated teeth, or bony
protuberances on their ns. These Paleozoic sharks could be a large as
14m, more than double the size of the current Great White Sharks.
.