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Origin and Evolution of Reptiles

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 Origin and Evolution of Reptiles

The origin of reptiles is well suited to maintain life on land. Many features show a considerable
advance in this respect over the Amphibia.

These are the:

(i) Dryness of the skin,

(ii) Reproduction, and

(iii) Devices for economising the use of water.

The origin of the reptiles lays about 320-310 million years back, in the steaming swamps of the
late Carboniferous, when the first reptiles evolved from advanced reptilomorph labyrinthodonts. The
oldest traces of reptiles is a series of footprints from the fossil strata of Nova Scotia, dated to 315
million years old. The tracks are attributed to Hylonomus, the oldest known reptile in the biological
sense of the word. It was a small, lizard-like animal, about 20 to 30 cm (8-12 inches) long, with
numerous sharp teeth indicating an insectivorous diet.

The early reptile Hylonomus

 Other examples include Westlothiana (for the moment considered to be more related to
amphibians than amniotes) and Paleothyris, both of similar build and presumably habit. One of the best
known early reptiles is Mesosaurus, a genus of early reptiles from the early Permian that had returned to
water, feeding on fish. The earliest reptiles were largely overshadowed by bigger labyrinthodont
amphibians such as Cochleosaurus, and remained a small, inconspicuous part of the fauna until after the
small ice age at the end of the Carboniferous.

EVOLUTION OF REPTILES

Reptiles evolved from amphibians of Carboniferous period, which depended on water bodies for
laying eggs and development of larval stages and hence could not exploit arid habitats far away from
water bodies. They invented a large yolk-laden shelled egg that could be laid on land and in which an
amniotic sac contained fluid in which embryo could develop to an advanced stage, capable of fending
for itself when hatched. The following anatomical changes transformed the ancestral amphibians into
land adapted reptiles:

1. Body developed a covering of epidermal scales to prevent loss of body moisture, and skin
glands were lost.
2. Skull became monocondylic for better movement and flexibility. Atlas and axis vertebrae
together permitted skull movement in all directions.
3. Limb bones and girdles became stronger but limbs were attached on the sides of body, and
belly touched the ground during creeping mode of locomotion.
4. Sacral region involved two strong and fused vertebrae to support the body weight on hind
legs.
5. Pentadactyle limbs developed claws that helped in climbing on rocks and trees.
6. Lung respiration became more efficient.
7. As a water conservation strategy, metanephros kidneys excreted uric acid which did not
require water for excretion.
8. Reptiles continued to be ectothermal since ventricle was not completely partitioned by a
septum and blood mixed in heart.
9. Internal fertilization evolved as a large cleioid shelled egg was laid on land.
10. Embryonic membranes amnion, allantois and yolk sac evolved to enable embryonic
development in arid conditions.

ANCESTORS OF REPTILES

I. PERMIAN REPTILES: THE COTYLOSAURS (ANAPSIDS)

The first reptiles are categorized as Anapsids, having a solid skull with holes only for nose, eyes,
spinal cord, etc. Turtles are believed by some to be surviving Anapsids, as they also share this skull
structure, but this point has become contentious lately, with some arguing that turtles reverted to this
primitive state in order to improve their armor. Both sides have strong evidence, and the conflict has yet
to be resolved. Cotylosaurs were the most primitive stem reptiles that evolved from the labyrithodont
amphibians (Embolomeri) in Carboniferous period. Seymoria was a lizard-like animal, with
pentadactyle limbs and a short tail. It had homodont labyrinthine teeth on the jaw bones as well as on
vomer and palatine bones. Presence of lateral line indicates its amphibious habits. Skull was
monocondylic for better movement of head. Seymoria indicates gradual transition from labyrinthodont
amphibians to reptiles. Another 5 foot long cotylosaur fossil, Limnoscelis was found in Mexico that had
large premaxillary teeth and long tail.
 With the close of the Carboniferous, reptiles became the dominant tetrapod fauna. While the
terrestrial reptilomorph labyrinthodonts still existed, the mammal-like reptiles evolved the first
terrestrial megafauna in the form of pelycosaurs like Edaphosaurus and the carnivorous Dimetrodon. In
the mid-Permian the climate turned dryer, resulting in a faunal turnover. The primitive pelycosaurs
where replaced by the more advanced therapsids.

The anapsid reptiles, with their massive skulls without postorbital holes, continued and
flourished throughout the Permian. The pareiasaurs reached giant proportions in the late Permian,
eventually disappearing at the close of the period (the turtles being possible survivors).

Early in the period, the diapside reptiles split into two lineages, the lepidosaurs (forefathers of
modern snakes, lizards, and tuataras). The group remained lizard-like and relatively small and
inconspicuous during the whole periode.

A = Anapsid, B = Synapsid, C = Diapsid

Very early after the first reptiles appeared, two branches split off. One lead to the Synapsida (the
"mammal-like reptiles" or "stem mammals"), having two openings in the skull roof behind the eyes
high, the other group, Diapsida, possessed a pair of holes in their skulls behind the eyes, along with a
second pair located higher on the skull. The function of the holes in bout groups was to lighten the skull
and give room for the jaw muscles to move, allowing for a more powerful bite. The diapsids and later
anapsids are classed as the "true reptiles", the Sauropsida.
II. THE PARAPSIDS

They possessed superior temporal vacuity in the skull and were adapted for aquatic mode of life.
Plesiosaurus was marine long-necked, fish-eating animal with 15 metre long fusiform body, short tail
and paddle-like limbs modified for swimming. The skull was euryapsid type with a superior temporal
vacuity. The fossils are from lower Jurassic (about 180 million years) and they are believed to have
become extinct in end-Cretaceous mass extinction.

Ichthyosaurus had fish-like body with fore limbs modified into paddle-like fins and hind limbs
disappeared. There was a fleshy dorsal fin too. Caudal fin was large and bilobed. Jaws projected into an
elongated snout and teeth were homodont, an adaptation for fish-catching. Skull was parapsid type with
additional postfrontal and supratemporal bones behind the eye orbit. Vertebral column became
secondarily simplified with amphicoelous vertebrae.
III. THE SYNAPSIDS
Synapsids split off from the primitive reptilian stock very early in evolution, perhaps in the
middle carboniferous period. Synapsids had started developing mammalian characteristics that enabled
them to be fleet-footed and active predators. Their legs commenced to move under the body. Heterodont
dentition and false palate started developing in pelycosaurs and had been completely formed in
therapsids. Two types of synapsids occurred from carboniferous to Permian, namely, the primitive
Pelycosaurs and advanced therapsids.

Pelycosaurs are represented by Dimetrodon whose fossils were discovered from North America and
Russia from the late Carboniferous to Permian periods. They were primitive reptile-like animals in
which limbs had moved under the body but not completely and each limb had 5 digits with claws.
Neural spines on the back were excessively long stretching highly vascularized skin between them that
formed a fin-like or sail-like structure. They had heterodont dentition with incisors, canines and molars
clearly defined but the false palate had not been completely formed.

Therapsids were more advanced and active synapsids which were perhaps endothermic animals with
high rate of metabolism. Heterodont dentition with false palate allowed these animals to chew and grind
food for quick digestion in the gut so that high metabolic demand of the body could be fulfilled. Jaw
muscles were attached to zygomatic arch to make chewing effective. Carnivore therapsids were called
Cynodonts (ex. Cynognathus) and herbivores were Dicynodonts.

IV. THE THECODONTS

They evolved from the sauropsid Archosauria, a group of insignificant lizard-like reptiles that
survived the Triassic mass extinction. They evolved into bipedal and highly agile predators. Euperkeria
and Ornithosuchus fossils were unearthed from South Africa and Europe. They were about 2 ft long
bipedal lizard-like animals with small head but very long tail for balancing while they chased flying
insects by rapid running. Endothermy must have evolved in thecodonts to meet the extraordinary energy
demands of their predatory life style.
V. THE SAURISCHIANS
They were dinosaurs with lizard-like pelvic girdle in which ischium and pubis bones radiated
away from each other. They were both bipedal and quadrupedal and carnivores as well as herbivores.

VI. THE ORNITHISCHIANS

They were dinosaurs with bird-like pelvic girdle in which ischium and pubis bones were directed
towards posterior as found in modern birds. These were also highly diversified carnivores as well as
herbivores
and both
bipedal and
quadruped.

VII. THE PTEROSAURIA

They were flying or gliding dinosaurs of Mesozoic that varied in size from sparrow-sized to
some species, like Pteranodon, having a wing span of 8 meters. They had pneumatic bones. Last digit of
the fore limb was extraordinarily long and served to attach the membranous patagium between fore
limb, hind limb and the body. Hind limbs were used for clinging on to the rocks and cliffs and 3 digits
of fore limbs also had curved claws, an adaptation for clinging. Their jaws were modified into beak that
possessed homodont dentition but Pteranodon did not have teeth.

Mesozoic, the "Age of Reptiles"

The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic extinction
event). Most of the earlier anapsid/synapsid megafauna disappeared, making room for the
archosauromorph diapsids. The archosaurs was characterized by elongated hind-legs and an erect pose,
the early forms looking somewhat like long legged crocodiles. The archosaurs became the dominant
group during the Triassic, developing into the well known dinosaurs and pterosaurs, as well as
crocodiles and phytosaurs. Some of the dinosaurs developed into the largest land animals ever to have
lived, making the Mesozoic popularly known as the "Age of Reptiles". The dinosaurs also deveoped
smaller forms, including the feather-bearing smaller theropds. In the mid Jurassic, these gave rise to the
first birds.

The lepidosauromorph diapsids may have been ancestral to the sea reptiles. These reptiles
developed into the sauropterygians in the early Triassic and the ichthyosaurs during the Middle Triassic.
The mosasaurs also evolved in the Mesozoic Era, emerging during the Cretaceous period.

The Therpasids came under increasing pressure from the dinosaurs in the early Mesozoic and
developed into increasingly smaller and more nocturnal forms, the first mammals being the only
survivors of the line by late Jurassic.

ORIGIN OF REPTILES

Reptiles have a dramatic evolution. They had an obscure beginning in the Palaeozoic, they
enjoyed a dominant status in the Mesozoic and then in the Coenozoic they faded out of scene.

I. Amphibian Origin:
It is generally accepted that primitive reptiles originated from some primitive labyrinthodont
Amphibia in the beginning of Carboniferous period. The emergence of such reptiles from labyrinthodont
Amphibia was so gradual that, on the basis of skeletal remains alone, it is difficult to ascertain whether
some of them are reptiles or Amphibia on way to transform to reptiles.
The labyrinthodonts possessed characteristically folded or labyrinthine teeth, similar to their
crossopterygian ancestors. They flourished through Carboniferous and Permian periods before
extinction in Triassic. We cannot, however, point out to a single ancestor of reptiles. Probably they arose
polyphyletically along a dozen or more independent lines.

II. Stem Reptiles (Cotylosauria):

During Carboniferous period of late Palaeozoic Era, about 250 million years ago, some
labyrinthodont amphibians gradually took on reptilian characters. These earliest reptiles are called the
stem reptiles. They belong to the order Cotylosauria of the subclass Anapsida. The transition was so
gradual that often it is difficult to decide whether some fossil skeletons are those of advanced
amphibians or primitive reptiles. They were contemporary of the primitive amphibians of late
Pennsylvanian and early Permian times. In the later part of the Permian, however, the amphibians were
outnumbered by these reptiles.
III. Ancestry through Seymouria:
One of the members of the Cotylosauria was Seymouria found in the Lower Permian sediments
in Texas (U.S.A.), perhaps 250 million years old. It was a lizard-like animal about 60 cm long, with a
comparatively thick body, relatively small pointed head with dorsally placed nostrils, and a short tail. It
had a few reptilian features like anapsid skull, single occipital condyle, large parietal eye and five digits.
Structure of Seymouria was intermediate between the amphibians of that time and the early reptiles.

Seymouria leads us to certain logical conclusions. It is not directly ancestral to all reptiles. At the
time it was living, the reptiles had already been present for some 50 million years. It is so perfectly
intermediate between an amphibian and a reptile that its true position remains uncertain. Romer treats it
as a reptile under the order Cotylosauria, whereas others classify it with primitive Amphibia under the
order Seymouria-morpha. Perhaps Seymouria is a connecting link between Labyrinthodontia and
Cotylosauria.

IV. Ancestry through Limnoscelis:

Limnoscelis was a genuine reptile from the late Carboniferous or early Permian of New Mexico.
Romer (1946) critically studied the Limnoscelis and suggested that Limnoscelis, a Captorhinomorph
Cotylosaur, is the primitive reptile. Limnoscelis was about 5 feet in length, some half of which was
made up by tail, with an elongated body, low slung on short stubby legs which sprawled outward from
the sides.
Like Seymouria, it was also aquatic in habitat. Its skull was of anapsid type and was compressed
from side to side and dorsoventrally flattened. The otic notch present in Labyrinthodonts had
disappeared leaving an indication at the back of the skull in the region of its closure.
The premaxillary teeth were enlarged and overhung the front teeth in the lower jaw, which is one
of the most common specialisation of early reptiles but seldom seen in amphibians. Other members of
the primitive reptiles are Captorhinus and Hylonomus.
V. Ancestry through Diadectes:
Diadectes was contemporary to Limnoscelis, but the didectomorphs represents different
evolutionary line from Limnoscelis. The didectomorphs retained the otic notch at the back of skull.
Diadectes had developed a specialised dentition. The front teeth were chisel-shaped and back teeth had
broad ridged crowns.
The first really large reptiles to make their appearance in the later Permian were the
Pareiasaurus, related to Diadectes. These lumbering animals ran to spiny excrescences on the head and
spiny armour of bony plates along back, which was presumably a defence against carnivorous mammals
like reptiles.
It is clear that the Cotylosaurs were, broadly speaking, ancestral to all reptiles. Some workers
including Watson believe that these Cotylosaurs separated into two divergent evolutionary lines. One of
these lines is represented by Captorhinomorph (suborder) of which Limnoscelis was primitive member,
led to mammal-like reptiles and ultimately to mammals and has been termed theropsida. The other line,
the Sauropsida, however, is represented by Didectomorph Cotylosaurs, the remaining reptiles and birds.
In this group the otic notch tends to persist.
Although, the authorities have not yet agreed, as at what precise stage of evolution these two
Theropsida and Sauropsida lines separated from one another. It seems clear, therefore, that the modem
reptiles are not intermediate in the evolutionary sense between amphibians and mammals. And to find
out a common ancestor for a creature like a lizard and a mammal, one has to go back to an ancient
reptilian or possibly to an amphibian, according to Watson.

Demise of the dinosaurs

The close of the Cretacious saw the demise of the Mesozoic reptilian. Of the large marine
reptiles, only the sea turtles are left, and of the dinosaurs, only the small feathered theropods survived in
the form of birds. The major surviving reptilian line is the lepidosaurs, of which the snakes are currently
the most numerous and widespread representatives. The end of the “Age of Reptiles”, opened up for the
“Age of Mammals”. Despite this, reptiles are still a major fauna component, particularly in tropical
climates. There are about 8200 extant species of reptiles (whereof almost half are snakes), compared to
5400 species of mammals (of which ⅔ are rodents and bats). The most numerous modern group with
reptilian roots are the birds, with over 9000 species.

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