COMPARATIVE EVOLUTION AND ANATOMY OF VERTEBRATE

All living organisms appear to be well-fitted or adapted for the problems posed by their respective
environment. It is easy to assume that either the characters of the organisms are fixed and have remained
so since the origin of life or that they are mutable and their diversity and adaptability have unfolded
progressively with time. This dynamic process is termed evolution. Evolution is defined as the gradual
and irreversible change from one condition to another over a very long period of time. Its principle
implies the development of an entity in the course of time through a gradual sequence of changes from
a simple to a more complex state. In this course, we will be looking at vertebrate evolution from the
simplest form which is the Jawless fish to the complex ones, the mammals.

Evolution of Fishes

The earliest and primitive fish, the Ostracoderms appeared in the Cambrian period about 530 million
years ago. They were without jaws, had primitive spinal columns called notochords and rudimentary
gills. An important change in this evolution is the use of gills for gaseous exchange which was used by
invertebrate chordates to filter food. They were also bilaterally symmetrical having two eyes, a mouth,
and a distinct tail. One of the primitive fish is Haikouichthys. In the Ordovician Period (480 million
years ago), the spinal column began to take on its modern form and the first true fish appeared in the
fossil record. Armored plates begin to develop on the head and thorax of fish. Astraspis was a jawless
fish covered with star-shaped scales from this time. Later, fish began to develop bony jaws and splits
into two distinct lineages the Placoderms and Acanthodians. Acanthodians were the first fishes with
jaws, the acanthodians, or spiny sharks were generally small shark-like fishes varying from toothless
filter-feeders to toothed predators. It is commonly believed that the acanthodians and the modern bony
fishes are related and that either the acanthodians gave rise to the modern bony fishes or that both groups
share a common ancestor. Placoderms were typically small, flattened bottom-dwellers. The upper jaw
was firmly fused to the skull, but there was a hinge joint between the skull and the bony plating of the
trunk region.

Osteichthyes were the early form of bony freshwater fishes until the Triassic. The Osteichthyes may
have arisen from the acanthodians. A subclass of the Osteichthyes, the ray-finned fishes (subclass
Actinopterygii), became and have remained the dominant group of fishes throughout the world. A major
extinction event occurred between the Triassic and Jurassic leading to the extinction of 70% of all
species of fish. During the Cretaceous Period and continuing into the Cenozoic Era we begin to see
more and more of the closest ancestors of modern fish such as Macropoma.

Adaptive features of fish

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1. Gills- Gills are special respiratory organs that absorb oxygen dissolved in water for respiration.
Gills helps the fishes to adapt to breathe or respire in water. They are present on either side of
the fish head. The gills are present in a gill chamber and are covered by a flap called
an operculum. The gills are involved in the gas exchange, ion
regulation, osmoregulation, hormone production, etc. In sardines, the gills perform a feeding
function too. The gills of fishes have a large surface area to volume ratio, enhancing gaseous
exchange.

2. Coloration- Fishes have a variety of colors and color patterns. Many fish have color patterns
that help them blend in with their environment thereby escaping predators. Such fishes include
the flatfishes, which change their skin colour to match the surrounding habitat. Fishes can also
have disruptive markings to hide body parts. Species such as the jackknife fish, high-hat and
some angel fishes, have dark lines that run through the eyes. These lines may serve to hide the
eyes so that other animals can not tell where the fish is looking or even if it is a fish. Also,
horizontal lines may be a sight-line for aiming attacks on prey. Fish coloration can also be
useful in catching prey. Many sharks exhibit coloration known as countershading. Sharks that
have countershading are dark on the dorsal side and light on the ventral side. With this color
scheme any prey looking down on the shark will see a dark shark against a dark sea bottom,
making it hard to detect the shark. Conversely, any prey looking up at the shark, will see the
light belly of the shark on the light background of the ocean surface water lit by the sun or
moon. Coloration can also be used for sexual attraction. An example is the darters and
sticklebacks, which use colour to attract and recognize potential mates.

3. Light organs- Some marine fish have the ability to produce light through bioluminescence.
Most light-producing fish live in mid-water or are bottom-dwelling deep sea species. In fish,
bioluminescence can occur two different ways: through symbiotic bacteria living on the fish or
through self-luminous cells called photophores. Some species of deep-sea angle fish may use
this light to attract prey, while others, like the Atlantic midshipman, may use this light to attract
mates.

4. Venom- Many fish may use venom as a form of defense. Most venomous fish deliver the toxins
through the use of a spine. Venomous spines are found in a wide variety of fish including
stingrays, chimaeras, scorpionfishes, catfishes, toadfishes, rabbit fishes, and stargazers.
Venomous spines can have poison glands along the grove of the spine, as with stingrays, or at
the base of the spine, as in some catfish. While humans can be stung by a multitude of fishes,
few species are life-threatening.

5. Electric organs- Elasmobranchs (sharks, skates, and rays) possess an electric sense system
known as the ampullae of Lorenzini. This system consists of many tiny gel-filled canals

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 positioned on the head of the fish. Through this system, these fishes are able to detect the weak
electric fields produced by prey. Some species of skates and rays also have electricity-
producing organs. The electric rays have paired electric organs located on either side of the
head, behind the eyes. With these organs, electric rays are able to shock and stun their prey.
Researchers believe that the skate’s electric organs are used for communication and mate
location. The electric eel can also produce electric fields. These eels use weak electric fields for
navigation, prey location, and communication. Additionally, these eels can produce strong
electric fields to stun potential prey. The strength of the “shock” is related to the size of the eel,
with larger individuals being able to produce more of a “shock.”

6. Sense Organ- Light does not penetrate to the deepest depths of the ocean, other senses have
become much more refined in fishes. Some fishes can detect vibrations in the water. A Shark
can detect one part per million of blood in the water.

7. Streamlined body- The streamlined bodies of fish are perfectly adapted to moving through
the water. While whales and dolphins are very distantly related to fish and evolved more
directly from land animals with little resemblance to fish, they feature similar body shapes.
This is an example of convergent evolution: the adaptive evolution of similar structures in
unrelated species to the same environment.

8. Fins and Tail- The presence of fins in fish aids in swimming. They are also used to maintain
the position of a fish during movement, and steering of fish in water. They also help in the
propulsion of the fish. Strong tails are present in fishes that act as a rudder to change the
direction and to keep their bodies balanced in water. Tails are also used to displace the water
so that the fish moves forward. Some differences in locomotion between fish highlight more
specific adaptations to individual ecologic niches. Forked or indented tails are seen in fish
that rely on swimming rapidly for long periods of time. Fish that do not travel extensively,
as part of their survival strategy, tend to have square or rounded tails, which are better
adapted to quick acceleration and stopping.

Evolution of Amphibians

Evidence from fossil records indicated that the first tetrapods: now-extinct amphibian species dated
back to nearly 400 million years ago. There is a major change in body plan from aquatic organisms that
respired and swam in the water, to terrestrial organisms that breathed air and moved onto land. These
changes occurred over a span of 50 million years during the Devonian period. Lobe-finned fish are
ancestral to amphibians. Their stump-like appendages and lung-like organs evolved into amphibian legs
and lungs. One of the earliest known tetrapods is from the genus Acanthostega. Acanthostega was an
aquatic organism having gills similar to fishes. However, it also had four limbs, with the skeletal

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structure of limbs found in present-day tetrapods, including amphibians. Therefore, it is thought that
Acanthostega lived in shallow waters and was an intermediate form between lobe-finned fishes and
early, fully terrestrial tetrapods. In 2006, researchers published news of their discovery of a fossil of a
“tetrapod-like fish,” Tiktaalik roseae, which seems to be an intermediate form between fishes having
fins and tetrapods having limbs. Tiktaalik probably lived in a shallow water environment about 375
million years ago. The early tetrapods that moved onto land had access to new nutrient sources and
relatively few predators. This led to the widespread distribution of tetrapods during the early
Carboniferous period: sometimes called the “Age of the Amphibians.”

In the Carboniferous period, the carnivorous amphibians started adapting to the terrestrial
environment and dominating them. Adapting to water conditions was different from adapting to land
conditions. Their lungs upgraded, and skeletons became more heavy and strong, making them better
at supporting their respective bodies’ weight on the land. They also gradually started
developing 'hands' and 'feet' and their skin had developed to retain body fluids and resist
desiccation. It was then that the amniotic egg started gradually developing and prevented the eggs from
drying and breaking out.

Adaptative features of Amphibians

Respiration: In the fishes, the gas exchange is carried out via the direct contact of water with the gills,
while in adult amphibians; the gas exchange is carried out through their moist and permeable skin. This
kind of respiration is called cutaneous respiration. Further, they also have lungs. During larval (tadpole)
stage, they respire only through gills and this is the reason that frogs and other amphibians need water
to survive.
Ecthotermy: Maintaining an appropriate body temperature is important for metabolism, locomotion,
and many cellular processes. Amphibians are ectotherms. Ectothermic organisms will often move to
new locations to increase or decrease their temperature if necessary. Some amphibians will find a
location in the sun and absorb sunlight to increase their temperature, this process is known as basking.
Amphibians in general have relatively moist skin, meaning that evaporative cooling occurs quickly in
the air. Amphibians maintain their temperature by moving locations or changing their postures.
Skin: Another adaptation in amphibians is their skin. Amphibians generally have soft moist skin without
scales. There are some exceptions. Toads have dry and warty skin. Skin acts as a preventive barrier
against changes in the external environment and can prevent cuts and microbial infections. The skin of
amphibians is unique because it can be used for gas exchange and respiration. In fact, some species
have lost lung organs over time. In addition to respiration, the skin has two types of glands present in
juveniles and adults: mucous and poison glands. Mucous glands secrete a protective coating over the

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skin. Some amphibian species have evolved poisonous secretions that serve as protection against
predators. Poisonous amphibians are typically brightly colored.

Evolution of Reptiles

The earliest amniotes evolved about 350 million years ago. They resembled small lizards, but they were
not yet reptiles. Their amniotic eggs allowed them to move away from water and become larger. They
soon became the most important land vertebrates. By about 320 million years ago, early amniotes had
diverged into two groups, called synapsids and sauropsids. Synapsids were amniotes that eventually
gave rise to mammals. Sauropsids were amniotes that evolved into reptiles, and birds. The two groups
of amniotes differed in their skulls. The earliest known reptile is Hylonomus. It was about 20 to 30
centimeters long, lived in swamps, and ate insects and other small invertebrates. At first, synapsids were
more successful than sauropsids. They became the most common vertebrates on land. However, during
the Permian mass extinction 245 million years ago, most synapsids went extinct. Their niches were
taken over by sauropsids, which had been relatively unimportant until then. This is called the Triassic
takeover. By the middle of the Triassic about 225 million years ago, sauropsids had evolved
into dinosaurs. Dinosaurs became increasingly important throughout the rest of the Mesozoic Era, as
they radiated to fill most terrestrial niches. This is why the Mesozoic Era is called the Age of the
Dinosaurs. During the next mass extinction, which occurred at the end of the Mesozoic Era, all of the
dinosaurs went extinct. Many other reptiles survived, however, and they eventually gave rise to modern
reptiles. The first turtle-like reptiles are thought to have evolved about 250 million years ago. Ancestral
crocodilians evolved at least 220 million years ago. Tuataras may have diverged from squamates
(snakes and lizards) not long after that. Finally, lizards and snakes went their separate ways about 150
million years ago. Reptiles separated from their water-dwelling ancestors and climbed onto land
during the Paleozoic era, over 280 million years ago. When that era gave way to the Mesozoic,
following a mass planetary extinction, reptiles survived and continued to evolve. They dominated the
earth between 248 and 213 million years ago and live on today as modern-day snakes, turtles, lizards,
and crocodiles.

Adaptive features in reptiles

Skin: Reptile skin contains keratin, a water-resistant substance that maintains hydration. Reptiles also
have scales to keep in moisture and help avoid skin damage, though the scales are sometimes too
small to be visible. This feature is most evident in turtles, whose scales fuse to form a shell.

Kidneys: Living on land means limited access to drinking water, so reptiles’ kidneys have adapted.
They conserve water by producing less urine in more concentrated forms.

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Reproduction: Laying soft-shelled eggs is safe in water, but land-dwelling creatures require a
different reproductive strategy. Scientists think this is why reptiles evolved a hard shell around their
eggs, and why some no longer lay eggs at all. In many types of snakes the eggs hatch internally, and
babies are born live. These shelled eggs, referred to as amniotic eggs, are an evolutionary adaptation
developed when amphibians were starting to evolve into the earliest species of reptiles. Unlike the
eggs of animals like frogs, which must be laid in water and defended because water offers no
protection to the growing embryo, shelled eggs wrap everything required for the embryo to grow
into a fluid-filled, protected package that can be laid on land and left alone.

Lungs: Adapting lungs in place of gills was a significant step in reptiles’ migration to land. While
amphibians all have gills at some stage in their development, either temporarily during the larval
stage or permanently through adulthood, reptiles are born with fully developed lungs.

Basking: For cold-blooded creatures on land, survival requires more than just physical changes. Since
a reptile’s temperature depends on its surroundings, it basks on rocks to warm its blood for hunting.
Without a place to bask, reptiles can’t get enough blood flow, as anyone who keeps reptiles as pets
can verify. Reptiles kept in captivity must have access to warming lights and heat-absorbent surfaces
to substitute for a natural basking environment.

Evolution of birds

Birds belong to a group of diapsids called the archosaurs, which includes three other groups: living
crocodilians, pterosaurs, and dinosaurs. Birds evolved within the clade Dinosauria, which is further
subdivided into two groups, the Saurischia and the Ornithischia. Saurischia diverged into two groups:
One included the long-necked herbivorous dinosaurs, such as Apatosaurus. The second group, bipedal
predators called Theropods, gave rise to birds. This course of evolution is highlighted by numerous
similarities between late (maniraptoran) theropod fossils and birds, specifically in the structure of the
hip and wrist bones, as well as the presence of the wishbone, formed by the fusion of the clavicles. A
well-known and important fossil of an animal that appears “intermediate” between dinosaurs and birds
is Archaeopteryx, which is from the Jurassic period. Archaeopteryx has characteristics of both
maniraptoran dinosaurs and modern birds. Some scientists propose classifying it as a bird, but others
prefer to classify it as a dinosaur. Traits in skeletons of Archaeopteryx like those of a dinosaur included
a jaw with teeth and a long bony tail. Like birds, it had feathers modified for flight, both on the forelimbs
and on the tail, a trait associated only with birds among modern animals. Fossils of older feathered
dinosaurs exist, but the feathers may not have had the characteristics of modern flight feathers.

Two basic hypotheses explain how flight may have evolved in birds: the arboreal (“tree”)
hypothesis and the terrestrial (“land”) hypothesis. The arboreal hypothesis postulates that tree-

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dwelling precursors to modern birds jumped from branch to branch using their feathers for gliding
before becoming fully capable of flapping flight. In contrast to this, the terrestrial hypothesis holds
that running (perhaps pursuing active prey such as small cursorial animals) was the stimulus for flight.
In this scenario, wings could be used to capture prey and were preadapted for balance and flapping
flight. Ostriches, which are large flightless birds, hold their wings out when they run, possibly for
balance. However, this condition may represent a behavioral relict of the clade of flying birds that were
their ancestors. It seems more likely that small feathered arboreal dinosaurs, were capable of gliding
(and flapping) from tree to tree and branch to branch, improving the chances of escaping enemies,
finding mates, and obtaining prey such as flying insects. This early flight behavior would have also
greatly increased the opportunity for species dispersal.

Figure 1: Evolutinary tree of birds and reptiles

Adaptative features in birds

Beaks: Varieties of beak shapes and sizes are an adaptation for the different types of foods that birds
eat. In general, thick, strong conical beaks are great at breaking tough seeds, and are found on seed-
eating birds such as cardinals, finches, and sparrows. Hooked beaks, such as those found on raptors like
hawks, eagles, falcons, and owls, are adept at tearing meat – perfect for these predatory birds. Straight
beaks of intermediate length are particularly versatile and are often found on omnivorous birds like
crows, ravens, jays, nutcrackers, and magpies. There are even highly specialized bills such as the

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flamingo’s: their beaks are comma-shaped for filter-feeding, enabling them to sift through mud and silt
in order to devour krill and other crustaceans.

Feet: The feet of birds have evolved as an adaptation to the landscapes they inhabit. Wading birds, such
as egrets and herons, have long toes to help with weight distribution as they make their way over reeds
and lily pads. Ducks and pelicans have webbed feet which aid in swimming. Some birds, such as the
American Coot, have lobate feet - a “halfway” point between webbed feet and long-toed waders to
assist in both modes of locomotion. Many bird species, like most songbirds, are also referred to as
“perching birds” because they have a foot structure that allows them to grasp branches - the
configuration of one toe at the back of the foot acts like a pincher, stabilizing the perched bird.

Nail: Nail structure plays an additional role in foot adaptation: the acute, strong nails of woodpeckers
and flickers give these species the ability to stand on and climb the vertical trunks of trees, a useful
adaptation for reaching insects that burrow beneath the bark. The grasping, sharp claws of a raptor, on
the other hand, are honed for subduing and even killing prey. Most running birds, such as ostriches and
emus, do not perch, therefore their back claw is either reduced or entirely absent.

Plumage: Plumage that is attractive to the opposite sex allows for more mating opportunities and, thus,
the ability to create more young. Additionally, feathers can disguise an organism, creating camouflage
for those that wish to hide from predators or sneak up on prey. The specialized flight feathers of owls,
on the other hand, are fringed for silent flight, making owls nearly impossible to detect as they swoop
down upon prey.

Evolution of mammals
Mammals evolved from a group of synapsids called therapsids. The synapsid lineage became distinct
from the sauropsid lineage in the late Carboniferous period, between 320 and 315 million years ago.
Following the Triassic takeover by Sauropsids, the surviving therapsids were small animals, some of
which evolved into cynodonts. During the first half of the Triassic Period, cynodonts thrived worldwide.
They were nocturnal and ate insects. They were saved from extinction mainly because of being
nocturnal. Animals in that niche were not taken over by the dinosaurs in the Triassic Period. As
Cynodonts continued to develop mammal-like traits, their evolution adapted them to their nocturnal
niche in several ways: They developed the ability to regulate body temperature which allows nocturnal
animals to remain active at night when the temperature falls. They also developed a good sense of
hearing because it is more useful than good vision if they are hunting in the dark. By the end of the
Triassic Period, cynodonts had shrunk in size and had many mammalian traits like a larger brain than
before, four different types of teeth, endothermy, three tiny bones in the middle ear, a diaphragm for
breathing, Lactation, and hair on the body. Although Cynodonts gave rise to mammals about 200

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million years ago, they are not considered mammals themselves. They became extinct during the
Jurassic or Cretaceous Period because of competition with early mammals.

Early mammals evolved from cynodonts and they continued to evolve. The first to split off were
Monotreme mammals, followed by marsupials. The last evolution of mammals was that of Placental
mammals. Early monotremes evolved about 150 million years ago. They retained some of the traits of
therapsids like laying eggs and having a cloaca. These traits are also found in modern monotremes.
Early marsupials evolved about 130 million years ago and one of the earliest of this kind was the extinct
genus Sinodelphys. It was about 15 centimeters (6 inches) long and its limb structure in the fossil found
hinted at it being a climbing animal. Climbing trees was a means for them to escape from predators by
climbing into trees. It survived on a diet of insects and worms. The ancestor of placental mammals may
have been the extinct genus Eomaia. Early placental mammals evolved about 110 million years ago and
fossils of Eomaia suggest that it may have had several traits of placental mammals. It may have been
only about 10 centimeters (4 inches) long and was a tree climber that ate insects and worms.
Evolution of Modern Mammals

The gradual and phased-out evolution of mammals led to the development of several features that
differentiated between modern mammals and modern reptiles. Several mammalian traits can be
attributed to their high levels of activity. A good example of this would be the system of double
circulation with a four-chambered heart, the diaphragm, anucleate and biconcave erythrocytes, and the
secondary palate that separates the food and air passages and facilitates breathing during suckling or
mastication. Hair on their body provides insulation which is known as endothermy or warm-
bloodedness. It helps in maintaining the body temperature of an individual independent of the
temperature of the external environment. Endothermy also allows mammals to have high levels of
sustained activity. The unique characteristics of mammals are a result of the evolution of their complex
and interrelated system.

Adaptative features in mammals

Adapting to their environment: Species of mammals have developed varying adaptations depending on
their environments. Mammals in cold climates have an insulating layer of either a thick coat of fur or a
thick layer of fat that help retain body heat and keep the animal's body temperature constant. Some
mammals that live in deserts survive by special adaptations in their kidneys and sweat glands, as well
as by their ability to avoid the heat by behavioral means. Other adaptations for survival in extreme
climates include hibernation or aestivation. These responses make it possible for the animal to

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conserve energy when food supplies become scarce.
Parental care: The care of their young ones is notable among mammals. Born at an average of 10% of
its mother's weight, mammalian young grow rapidly. The protection the young receive from one or both
parents during the early stages of their lives enables mammals to maintain a strong survival rate in the
animal kingdom. The subclass Placentalia contains the majority of living mammals. The embryo of
placentals develops in the mother's uterus, is nourished by blood from the placenta, and is retained until
it reaches an advanced state of development. The Marsupialia are found in Australia and in North
and South America. Their young develop inside the uterus of the mother, usually with a placenta
connected to a yolk sac. Young marsupials are born in a very undeveloped state and are sheltered in a
pouch (the marsupium) which contains the nipples of the milk glands. Kangaroos, wallabies, and most
Australian mammals are marsupials, as is the opossum of the New World. The Monotremata of
Australia includes the duck-billed platypus and two species of spiny anteaters. Monotremes lay eggs,
but have hair and secrete milk like other mammals.

Endothermy: Endothermy comes from Greek root words meaning heat within. Endothermic animals
are sometimes described as being warm-blooded. All mammals are endothermic and use a variety of
mechanisms to maintain steady, homeostatic internal body temperatures. Homeostasis is the condition
of a body system that is actively regulated to remain consistent. If a mammal’s body temperature begins
to fall, it can shiver or increase its metabolic rate of converting food energy to heat. If a mammal begins
to overheat, it can secrete sweat or increase blood flow to the skin to cool off. These mechanisms allow
mammals to thrive in a wide range of environments.

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